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LIFE. Illustrated. 










Director of the Scripps Institution for 

Biological Research of the University 

of California, La Jolla 








All Rights Reserved 

This work contains the text of the book: 
'An Organismal Theory of Consciousness.' 

Made in the United States of America 

The Gorham Press, Boston, U. S. A. 





B. The Production of Individuals by Other Indiridiuils (Concluded) 



HEREDITY .......... 1 

Evidence from spermatozoa, 1. Spermatozoa subject to 
heredity as well as "bearers of heredity," 1: (a) Illus- 
trated by the ontogeny of mammalian sperm, 4; (k) Illus- 
trated by the ontogeny of an insect sperm, 9. Evidence from 
the ovum, 15: (a) eggs of ascidians the facts, 16; (6) 
Conklin's interpretation, 19. Critical examination of Conk- 
lin's interpretation, 23. 



The mitochondrial theory of heredity, 32. The mitochon- 
dria! theory tested by the ontogeny of spermatozoa, 36. The 
mitochondrial theory tested by histogenesis, 37. The un- 
tenable hypothesis that cytoplasm of the ovum is inheritance 
material for general but not for special characters, 40. Spe- 
cies attributes in single cells of adult organisms, 43- The 
spinules of the ascidian genus styela, 44- The spicules of 
sponges and other invertebrates, 50. The '"hairs" of higher 
plants, 55. Cell-wall structures in higher plants, 57. The 
morphology of striated muscle fibers, 60. The physiology of 
muscle fibers, 61. Summary of positive information about 
the physical basis of heredity, 64. 



Antecedents of the cytoplasmic and nuclear theories of in- 


x Contents 


heritance material, ('>(]. Function of chromosomes in heredity 
acquired and secondary, 67. The tiro-fold character of the 
problem of hereditary substance, 70. The probability that 
inheritance material becomes snch in each ontogeny, 73. 
Germ-cells subject to metabolism like all other cells, 74. 
Chemical changes in germ-cells during parent's ontogeny, 75. 
The possibility of changing sex by influences on the germ, 
70. The determiner conception contrary to ordinary chem- 
ical principles, 79. Endorsement of E. B. Wilson's proposal 
to drop "determiner" from the vocabulary of genetics, 82. 
Advantages in conceiving germ-cell chromosomes as initia- 
tors in hereditary development, 83. Inconclusiveness of the 
cy to-logical evidence usually appealed to in support of the 
chromosome theory, Xj. Humming up of the findings 
against the chromosome theory, 87. Brief reference to the 
untoward implications of the germ-plasm conception of 
heredity, 89. 



The field to be covered by the constructive discussion, 93. 
Four types of bio-integration to be treated, 9^. Graded 
repetitive series as integrative phenomena, 95. Illustrations 
from, animals, 95. Illustrations from plants, 99. Justifica- 
tion for bringing all these phenomena under one head^ 103. 
Attempted casual explanation of these series, 104. Asial 
metabolic gradients as integrative phenomena, 107. M eristic 
gradients and metabolic gradients both phenomena of growth 
integration, 111. 



Functional as contrasted with growth integration, 113. The 
conception of "internal secretions," ll//. Effects of removing 
the human thyroid for curative purposes, 115. Experimental 
thyroid excision in normal lower animals, 117. The internal 
secretion of the duodenal mucous membrane, 119. The na- 
ture of the active substances in internal secretions, 121. The 
clone resemblances and interrelations of the different in- 
ternal secretions, 12^. Relation between the internal secre- 

Contents xi 


ion/ and nerrous systems, 128. Composition and nature 
of (he autonomic system, !..",>. Experimental evidence of 
connection between the adrenal gl-ands and the nervous sys- 
tem, 1.11. Clinical evidence of adrenal-nervous connection, 
133. Sum mar if of present state of knowledge in this field, 



Genera] inability of elementalism to interpret the phenom- 
ena, 1'fl. Cri/!<iue of the view that internal secretions are 
'''formatire stuff's," l.j?. The meta/thysical and logical weak- 
ness of the riew, 1/jX. The form of metaphysical absolutism 
involved, lol. Confusion of theory of organisms and 
theory of the knowledge of organism*. 1~>2. An illustra- 
tion of neglect of fact by elemenlalist theory, 1~>7. A peculiar 
elementalist objection to the organic whole, 13S. 


Distinction, between developmental and functional integra- 
tion, 161. Neural and not psychical phenomena the subject 
of this chapter, 161. The authors indebtedness to Sher- 
ring ton's work, 16.'. The fundamentally of cellular inte- 
gration in the re-flex-arc, 163. The integration of reflex- 
arcx. }>;s. The spreading and compounding of reflexes, 171. 
Antagonistic re-flexes in skeletal muscle groups finally in- 
tegral ire, // '/. The antagonisms within the autonomic sys- 
tem finally integratire, 178. Concluding remarks on the sig- 
nificance of neural integration for the organismal standpoint, 


XEHVE ACTION ......... 185 

Neglect of the works of Sherrington and Cannon by Jacques 
Loeb, 18o. The real importance of Loeb's conception of the 
nerrous system, 7<s'/\ The organismal character of tropisms 
partly recognized by Loeb. 188. The organismal character 
of the segmental theory of the nerrous system, 190. Critique 
of the elementalist attempt to interpret tropic-tic and seg- 
mental theories of the function of the nerrous system, 19S. 
The bearing of this critique on "analysis" in Biological 

xii Contents 


reasoning, 206. Theories of animal behavior in relation to the 
"science" of zoology, 208. 


Preliminary remarks: (a) Absolute discrimination between 
reflex and psychical phenomena not necessary, 214 j (&) The 
organism an original datum in all problems of psychic life, 
215; (c) Provisional classification of psychical facts, 217. 
Likeness between tropistic and higher psychic activity, 220. 
First move toward showing the organismal character of the 
higher psychic life, 227. Associationist psychology a special 
case of elementalist biology, 228. Preliminary examination 
of objective and subjective, 231. The essence of Wundtian 
apperception, 232. Remarks on analysis and synthesis, 236. 


A still closer look at the organismal nature of tropisms, 
239. The automatic and anticipatory character of tropisms 
and .other reflexes, 2^1. A still closer look at the likeness 
between higher rational life and tropisms, 242. A still 
closer description of the subrational moiety of psychic life, 
246. Remarks on the classes of subrational life, 246- Four 
certainties about the adaptiveness of subrational psychic 
activities, 250. Generally useful to individual and to species, 
250. Many useful to species primarily, 250. Variability of 
subrational activities, 251. Tendency of subrational activ- 
ities to e.vcessiveness, 256. Summary of organismal charac- 
ter of all sub rational psychic- life, 274- Specif city of subra- 
tional psychic life, 276. 

Remarks on the hypothetical character of this chapter, 282. 

The natural history method and the study of one's self, 282. 
Formulation, of the central hypothesis, 286. Preliminary 
justification of the hypothesis as such, 287. More systematic 
justification of the hypothesis, 291. The nature of "outei-" 
or objective and "inner" or subjective, 292. As to the lowest 
terms of self -consciousness, 308. Instinct and physical or- 
f/anhiition, 310. Emotion and physical organization, 316. 
Glance at the eqnillb rathe interaction between "body" and 
"soul," 323. Support of the hypothesis by the physico-chem- 


Contents xiii 


if-al conception of the organism, 324- Personality and ele- 
mentary chemical substances, 327. On the psychology of 
subjective and objective personality, 331. Personality and the 
"breath of life" viewed in the light of physical chemistry 
of the organism, 336. 


BIBLIOGRAPHY ........... 359 

GLOSSARY ............ 377 




56. Skeleton of Python Frontispiece 

36. Sperm of Fur Seal (After Oliver) 4 

37. "Young" Spermatid (After Ballowitz) 5 

38. Spermatid of Fur Seal (After Oliver) 5 

39. Spermatid of Fur Seal (After Oliver) 6 

40. Spermatid of Fur Seal (After Oliver) 8 

41. Spermatid of Fur Seal (After Oliver) . . . . . 8 

42. Development of Sperm of Argas Miniatus (After Casteel) 10, 11 

43. Development of an Ascidian Egg (After Conklin) ... 17 

44. Spinule Cell of Styela Yakutatensis (After Huntsman) . 44 

45. Spinule Cell of Styela Plicata (After Huntsman) ... 45 

46. Spinule Cell of Styela Greeleyi (After Huntsman) . . 46 

47. Spinule Cell of Styela Montereyensis (After Ritter and For- 

syth) 47 

48. Spicules of Sponges (After Lankester) 51 

49. 50, 51. Development of a Spicule (After Lankester) . 52, 53 

52. Trichomes of Papaver Orientale (After Cannon) ... 55 

53. Trichomes of P. Pilosum (After Cannon) .... 55 

54. Trichomes of P. Somniferum (After Cannon) .... 55 

55. Side View of Auiphioxus (After Parker & Haswell) . . 95 

57. Tentacle of Halocynthia Johnsoni (Schematic; After Ritter) 98 

58. Acacia Elata 99 

59. Vicia Gigantea 100 

60. Cassia Sp 100 

61. Sequoia Sempervirens 101 





B. The Production of Individuals by Other Individuals 


Chapter XIV 


Evidence from Spermatozoa 

f N our discussion we will be guided by the principle laid 
down earlier and followed throughout our treatment 
of heredity in the Protozoa, namely, that descriptive onto- 
genesis brought to bear on the actual transformations which 
result in the production of specific organs and parts, is the 
final tribunal for the determination of what substances are 
hereditarily formative. The first inquiry will be whether 
there exist among the metazoa single-cell organs or parts 
which when fully formed exhibit species characters in the 
sense of taxonomic biology. If such elements do exist, ob- 
servation on the constituents of the undifFerentiated cells 
which take part in the transformations, obviously may be 
expected to give us the information sought as to what 
substances are formative. 

Spermatozoa Subject to Heredity as Well 'as "Bearers of 


The comparative anatomy and comparative ontogenesis 
of the male germ-cells among animals, which have been pur- 
sued with great assiduity and skill during recent decades, 


2 The Unity of the Organism 

furnish perhaps the largest mass of relevant facts we pos- 
sess from any one field. 

Innumerable researches on fully formed spermatozoa, the 
greatest single research being that of Retzius, give us ex- 
tensive knowledge of the variety of structure of the sperma- 
tozoa in the larger and smaller taxonomic subdivisions of 
the animal world. It would be going beyond the evidence 
to say that every well-characterized animal species may be 
identified by its spermatozoa; but unquestionably the trend 
of investigation has been toward such a conclusion. I 
believe, for example, it would be impossible to assert on the 
basis of evidence that any two species of animals belonging 
to different genera, no matter how much alike if their dis- 
tinctiveness is not questioned, have indistinguishable sper- 
matozoa. "One may say," writes Ballowitz, "that each ani- 
mal species has its own sperm-form of definite size." An 
attempt to illustrate fully this variety of form and size by 
specific examples is out of the question here. We will refer 
only to the specificity of the sperm of man himself. Ret- 
zius was able to compare in detail sperms of the Chimpanzee, 
Orang-Utan, Gibbon, and Homo, and found that while their 
resemblance is rather close, each possesses clear differential 
marks. For example, the spiral structure of the envelop of 
the central piece is considerably more distinct in the Chim- 
panzee than in Homo. Worthy of mention is the fact that, 
according to Retzius, the sperm of the Chimpanzee resembles 
that of Homo more closely than does that of the Orang, 
thus falling in with the fact that in several particulars of 
adult structure the resemblance of the Chimpanzee to man 
is closer than that of the Orang. 

The spermatozoa of a given animal group having a closer 
resemblance to one another than to those of other groups ; 
in other words, having a resemblance due to descent, are 
themselves subject to heredity and are not alone concerned 
in the transmission of hereditary attributes from parent to 

Evidence from. Metazoan Germ-Cells 3 


Especially important for us is it to notice that to a great 
extent the diversity of structure among spermatozoa is in 
the locomotor organ, the tail; that is, the organ chiefly 
concerned with the unique life of the sperm as such, and 
very little if at all concerned directly with fertilization and 
hence with hereditary marks of offspring. This fact de- 
serves attentive consideration. "In its more usual form the 
animal spermatozoon resembles a minute, elongated tad- 
pole, which swims very actively about by the vibration of a 
long, slender tail." 2 In some respects comparison of the 
spermatozoon of the type here indicated with an Appen- 
dicularian, a minute Tunicate which possesses a tail through- 
out its life, is more instructive. Any one who has had op- 
portunity to observe both sperm cell and appendicularian 
when alive and active will not have failed to remark the 
general resemblance, not only as to form but as to kind 
of movement in the two cases. Is the development of the 
Appendicularian's tail a manifestation of heredity? Surely 
no one would think of giving any but an affirmative answer. 
How, then, deny that the development of the spermatozoon's 
tail is also a manifestation of heredity? I cannot see that 


it would be less inconsistent to affirm that the wriggling ap- 
pendicularian is alive but that the wriggling spermatozoon 
is not, than to affirm that the ontogeny of the first is guided 
by heredity while that of the second is not. To bring the 
point onto somewhat more familiar ground, let us revert 
to Wilson's comparison of the spermatozoon to the tadpole 
stage in the life of the frog. Our contention is that the tail 
of the frog's spermatozoon is as indisputably modeled by 
heredity as is the tail of the frog's tadpole, and consequently 
that we are bound to search for the physical basis of hered- 
ity in the former as well as in the latter. Our inquiry is, 
then, What observations have we as to the substances con- 
cerned in producing the spermatozoon tail? 

The Unity of the Organism 

(a) Illustrated by the Ontogeny of Mammalian Sperm 

Without exception, so far as I know, positive description 
of spermatogenesis affirms that the tail is produced from 




h.c., head cap. n'k., neck, c.p., connecting piece, c.r., cytoplasmic 
remnant, g.a., anterior granules. g.p., posterior granules, an., 
aimulus. m.p., main piece, e.p., end piece. 

other parts of the spermatid than the nucleus. For the lit- 
tle we can do in the way of giving objectivity to this general 
statement we will first examine the nearly mature, typical 

Evidence from Metazoan Germ-Cells 5 

mammalian spermatozoon (figure 36) of the Fur Seal. 

For the origin of the various parts we will make use of 
the summary given by Ballowitz (figure 37). Concerning 



1, nucleus. 2, centrioles. 3, icliosome. 4, mitochondrial body. 5, 
chromatoid body. 6, spindle remnant. 

the nucleus, 1, "it is certain that in all animals the chroma- 
tin-containing part of the sperm proceeds from the 
nucleus." 3 On this point there is agreement among observ- 
ers ; so whatever may be the truth about the sperm-chromatin 


n., nucleus, c., centrioles. a.f., axial filament. 

as the physical basis of heredity for the adult animal, there 
is no question about its being such for the head of the 
spermatozoon itself. 


The Unity of the Organism 

The centrioles (2, figure 37) are "almost always double and 
occupy a place in the spermatid close beneath the surface of the 
cell." 3 The pair is typically so placed that the axis joining them 
is perpendicular to the surface of the spermatid, the member 


h.c., head cap. n., nucleus, p.c., proximate centriole. 
centriole. a.f., axial filament, s., remnant of sphere. 

d.c., distal 

toward the surface being known as the distal centriole and the 
one toward the center of the spermatid the proximal centriole 
(figure 39 d.c. and p.c.}. 

As to the part played by each of these in the development of 


n., nucleus, c.h., chromatin granules, c.t.f., caudal tube filaments. 

the sperm we learn that after migrating inward until they 
come to lie very near the nucleus, if not in actual contact with 
it, the centrioles begin their development with the following out- 
come: "The proximal centriole . . . divides into two portions, 
closely adherent to the nuclear wall, each connected by a fila- 
ment to one distal group. The distal centriole divides into an 

Evidence from Metazoan Germ-Cells 7 

anterior and a posterior portion. The posterior portion becomes 
the annulus (an., figure 36) while the anterior one divides again, 
forming the Xoduli posteriores" 4 (g>p-, figure 36). Throughout 
their career these bodies or granules are highly stainable with 
certain dye-stuffs. 

Out of the idiosome (3, figure 37) which as a rule is a body 
"fur sich" -in itself alone develops the perforatorium, or head 
cap of the adult sperm (h.c., figure 36). It is agreed that the 
mitochondria (4, figure 37) in the spermatids of many animals, 
particularly of many vertebrates "furnish the material" for the 
spiral found in the connecting piece (c.p., figure 36) of the 
sperm. Although no spiral is present in the Seal sperm it may 
be represented, according to Oliver, by numerous granules sur- 
rounding the axial filament in the connecting piece. But while 
the connecting piece of the Seal sperm seems not to be typical 
as regards the spiral, it presents another structure, the caudal 
tube, or "manchette" of some authors, in a form which is specially 
instructive from our standpoint. In the adult sperm this struc- 
ture is a thin sheath enveloping the cytoplasmic part of the con- 
necting piece and lying in close contact with the persisting cell 
membrane. The point of special interest about it is that its 
persistence in the completed sperm of the Seal appears to be 
exceptional, for it is known to disappear entirely in the course 
of development of the sperm of several other mammals. It is a 
transitory or embryonic organ in some species of sperm, but a 
permanent one in other species, just as gills, for example, are 
transitory organs in the ontogeny of some species, as a frog, but 
are permanent in others, as fish. 

The development of the tube in the Seal sperm is especially 
favorable for observation. "It may be readily followed," writes 

V V 

Oliver, "from its first appearance up to its final incorporation 
in the connecting piece as a peripheral layer, or sheath." 4 Here 
then is a structure having all the essential marks of devel- 
opment due to heredity and likewise one the "physical basis" of 
which has been carefully observed. Mentioning a long list of 
investigators who believe in the "derivation of the caudal tube 
by a process of cytoplasmic differentiation alone" Miss Oliver 
tells us that her study of the development of the fur seal sperm 
is a complete confirmation of this view. As to the very begin- 
ning of the tube we read: "Shortly after the centrosomes and 
their tail filament have reached the nuclear membrane there 
appears in the cytoplasm surrounding the axial thread a series 


The Unity of tlie Organism 

of delicate filaments attached to the nuclear membrane. The 
proximal ends of these arise in a circle around the basal end 
of the nucleus with the centrosomes as a center, while their 
distal ends project freely into the cytoplasm." 5 (figure 4-0 c.t.f.} 
These filaments "are at first very short and thin, but they in- 
crease in length and thickness rapidly. By the progressive dif- 
ferentiation of the cytoplasm between them they soon fuse into 
a hyaline tube, surrounding the axial thread and open at its lower 
extremity." (figure 41, c.t.) The capital point is that we have 
here a well-defined structure the development of which is in- 



h.c., head cap. c.t., caudal tube, an., annulus. 

dubitably proved to depend primarily on parts of the cell other 
than the chromatin. Indeed no one, apparently, has pretended 
that the chromatin takes a part in its production, for even those 
investigators who have not believed that it arises from the cyto- 
plasm alone have held that it originates from the membrane of 
the nucleus. 

About the chromatoid body (-5) and the "spindle remnant" (6} 
(figure 37), little need be said in this connection as they seem to 
be inconstant structures the significance of which is in much 
doubt. Finally, mention should be made of the fact that a con- 
siderable portion of the cytoplasm is cast off entirely in the 
ontogeny of the sperm of many animals, as for example the seal, 

Evidence from Metazoan Germ-Cells 9 

the body (c. r., figure 36) being all that is left of the substance 
at the late stage represented. So much by way of illustration of 
the portions of the spermatid, or germ of the spermatozoon, in 
the vertebrated animals, for despite the great variety in struct- 
ural details presented by the sperm of this part of the animal 
kingdom I think all will agree that so far as concerns the chief 
point being made in this discussion, what we have presented is 
true of the whole phylum. 

(b) Illustrated by the Ontogeny of an Insect Sperm 

We will now examine the ontogeny of a very different 
type of sperm, from another portion of the animal kingdom, 
the insecta. The particular species chosen is the fowl-tick 
(Argas miniatus). The investigation made use of is by 
Doctor D. B. Casteel. 

The series of figures (42 a, b, c, d, e, f, g,) will help to 
an understanding of the remarkable, almost unique sperm and 
spermogenesis in this animal. Figure 42a shows the nearly ma- 
ture primary spermatocyte. Especially to be noted are the 
mitochondria, mi., scattered uniformly through the cytoplasm, 
and the striated layer, s. L, on the outer surface of the cell. 
This layer is sharply demarked from the underlying cytoplasm. 
The striae, disposed perpendicular to the surface of the cell, are 
excessively fine, and when looked at in situ end on "suggest the 
appearance of a faceted compound eye or of honey-comb." 6 
Concerning the genesis of this layer Casteel says, in a personal 
letter, that the layer begins to appear at the surface and grad- 
ually increases in depth until the completed state shown in figure 
42a is reached. 'The striae," he says, "appear to be forming 
from the undifferentiated cytoplasm sheath." 

All the figures from 42 b to g have to do with the transforma- 
tion of the spermatid (figure 42b) into the spermatozoon. From 
figure 42b one sees that the striated layer has disappeared on one 
side of the cell and thinned out greatly in a smaller area on the 
opposite side; that the nucleus, n, has moved to the surface of 
the cell in the middle of the area of disappearance of the striated 
layer; that the large plasmosomes, pi. 42a have almost entirely 
disappeared from the nucleus; that the vesicular bodies v.b., as- 
sembled for the most part in the vicinity of the nucleus, are 

q " 



a., ru})ture point of outer tube, c.p., cilia-like processes, fl., flagel- 
lum. f.p., finger-form process, g.e., gelatinous envelop, i.e., in- 


:x-- St. 




_ \. m 


vagination cavity, i.t., inner tube, mi., mitochondria, m.r., mito- 
chondrial ring, n., nucleus, o.d., oil droplet, o.t., outer tube, pi., 
plasmosome. s.l., striated layer, v.b., vesicular bodies, b.p., begin- 
ning of inner tube. 


The Unity of the Organism 

in processes of degeneration and disappearance; and that the 
mitochondria, mi, look as though they had collected into two 
well-defined spherical masses m.r., on the side of the cell oppo- 
site the nucleus and near the small area of thinned-out striated 
layer. As a matter of fact these two apparent mitochondrial 
masses are the opposite sides of a ring seen in optical section. 

Transformation of the general form of the spermatid now be- 
gins by the indentation of the side opposite the nucleus, this 
going on to produce first the quarter-moon shape shown at i.e., 
figure 42c. By the still further growth and narrowing the edges 
of the cup finally come together to produce the elongated cavity 
shown in figure 42d, o.t. This is the beginning of the outer tube 
which becomes long and relatively narrow as development con- 
tinues, (figure 42e, o.t.}. At an early period in the growth of 
this tube the striated layer which naturally becomes shut into the 
tube breaks up over most of the circumference of the tube into 
what resembles a dense layer of long cilia. However,, since the 
processes are not motile and later dissolve and produce a gela- 
tinous mass within the tube, their resemblance to cilia is only 
superficial. This account of the fate of the striated layer applies, 
as previously intimated, to most of the circumference of the outer 
tube. But the small thin area of the layer opposite the nucleus, 
shown in figures 42d and 42c, retains the cilia-like processes. 
This persistent basal patch (b.p., figure 42d) is the starting 
point for an important part of the future spermatozoon, the 
"inner tube" so called by Doctor Casteel (i.t., figure 42e). 

While these profound changes are going on in the portion 
of the spermatid opposite the nucleus, a stout, somewhat finger- 
like process, (f.p., figure 42d) is formed on the nuclear side of 
the spermatid, into the base of which the nucleus migrates. In 
the meantime the vesicular bodies have entirely disappeared, and 
the mitochondria, no longer disposed in the ring of earlier stages, 
have assembled into an irregular, rather diffuse mass toward the 
basal patch (mi., figure 42d). 

At the time when the outer tube has reached its maximum 
length and is somewhat coiled, the inner tube, starting at the 
basal patch previously described, begins to grow into the cavity 
of the outer tube. This growth continues until the inner tube 
is approximately as long as the outer tube. Figure 42e presents 
an advanced but not completed stage of growth of the inner tube 
i.t. In reality, according to Doctor Casteel's interpretation, the 
inner tube grows at the expense of the outer tube, for when the 

Evidence from Metazoan Germ-Cells 

two are of nearly equal length the entire spermatozoon is only 
about half as long as it was before the inner tube developed. 
All of the mitochondria of the cell are drawn into the inner tube 
as it grows, and finally form a deeply staining mass at the distal 
end of the tube (mi., figures 42e and 42g). 

During these transformatory operations the nucleus,, greatly 
reduced in size proportionally to the spermatozoon as a whole, 
has left its former place at the base of the finger-like process, 
and been making its way along the wall of the outer tube, bur- 
rowing through the gelatinous layer on the outer part of this 
tube (n, figure 42e). This migration continues until, when the 
distal end of the inner tube reaches nearly the end of the outer 
tube, the nucleus lies in the wall of the outer tube opposite the 
end of the inner tube (n, figure 42f.) 

The final act of transformation takes place after the sperm 
has left the male tick and lies in a spermatophore sac within 
the genital ducts of the female. This act begins with the per- 
foration of the end (a. figure 42f) of the outer tube by the 
inner tube. Through the opening thus made the whole inner tube 
finally passes, really by a slipping back of the outer tube, so 
that, the eversion of this latter being completed, the two tubes 
constitute one continuous tube. By this act of turning inside 
out, the finger-like process in which the nucleus formerly lay 
(figure 42d, f.p.} is brought into close proximity again with the 

Some of the details of the final steps in the transformation 
Doctor Casteel has not yet been able to make out; but these 
are of little consequence for our discussion. Nor has the 
act of fertilization been observed. Going on the usual cri- 
teria, the end of the sperm containing the nucleus would be 
regarded as the head. But surprisingly enough, in moving, 
the opposite end, the end containing the mitochondria, goes 

This sperm and its development are so unique as con- 
trasted with those occurring in most animal groups, that 
one might be almost inclined to question whether there may 
not be something wrong here whether the case may not be 
one of diseased growth, or the result of manipulative mal- 

The Unity of the Organism 

formation, or something else. Any such suspicion is, how- 
ever, completely done away with by the fact that much the 
same type of spermogenesis is known to occur in other ticks. 
Casteel cites particularly the observations of Katharine 
Samson on the development of the sperm of Ixodes ricinus 
and Ornithodes moubata as furnishing cases to which that 
of Argas is "in many respects parallel." 

After all that has been said in the previous pages, it is 
almost needless to point out the significance for our general 
contention of this remarkable case of spermogenesis. Were 
we living in that comfortable era of life-philosophy wherein 
theologians studied nature for the purpose of proving that 
everything in it was made to meet some human need, we 
could easily recognize this case as one designed expressly 
to assist man in refuting the false dogma of Chromatinic 
Omnipotence in heredity. 

It is hard to imagine a developmental process in which 
denial of form-determining power to non-chromatinic, even 
non-nuclear parts of the cell would be a greater folly than 
would be the denial of cytoplasmic "form-determination" 
in the production of the striated layer of the spermatid, or 
of the growth of the outer tube, or of the inner tube, or 
of the turning inside-out of the outer tube. In fact, by far 
the greater part of the astonishing transformations here 
gone through are cytoplasmic, the nuclear changes being 
relatively slight. 

It may be worth while to remind the reader again, so 
boldly does the real truth about hereditary substance stand 
out in this case, that the cardinal evil in the chromatin 
dogma is that it implies the denial of great masses of the 
most direct observational evidence we have as to what the 
physical bases of heredity may be, and so tends to detract 
attention from them. We may predict that the important 
research which has made known this unique case of organic 
genesis will pass almost unnoticed by the geneticists of our 

Evidence -from Metazoan Germ-Cells 15 

day. Were their attention called to it they would probably 
frankly say that they have little interest in genesis in this 
sense. That the sperm here described is not peculiar in 
every respect to the species Argas miniatus is certain from 
the meager comparative information we possess, as Casteel 
has shown. Nevertheless not merely general analogy, but 
strong indications contained in even the little comparative 
knowledge we have in this particular case, warrant the sup- 
position that in some respects the sperm of the species would 
be peculiar to the species, to say nothing of the genus, 
family and so forth. The development is, consequently, due 
to heredity, and the cytoplasm is "inheritance material" 
as ascertained by direct observation. 

Evidence from the Ovum 

We now turn to the ovum to see what can be learned con- 
cerning hereditary substance in the development of the 
ovum iself. Attention should be called at the outset to the 
important difference between the sperm and the ovum in 
the kind of specialization in each. The sperm, it will be 
noticed, is far more specialized for its own particular life 
than is the ovum, this "particular life" of the sperm con- 
sisting in its great power of locomotion. As a consequence 
of this difference, the ovum as an entity has no such sharp 
distinction from the ovum as a germinal element as has the 
spermatozoon. This difference is expressed in one way 
by the assertion that the fertilized ovum is the individual 
organism in the one-celled stage of its life. No such state- 
ment is ever heard about the spermatozoon for the obvious 
reason that the sperm does not transform directly into the 
embryo as does the egg. From the absence of so distinctive 
a character of the ovum as such, it happens that the hered- 
ity of the ovum is not so distinguishable from the heredity 
of the organism of whose life it is a stage, as is the case 

16 The Unity of the Organism 

with the sperm. Nevertheless we are bound to recognize 
that the egg no less than the sperm has hereditary attri- 
butes of its own, and that other substances than chromatin 
play a demonstrable part in the production of these. In- 
deed the main discoveries concerning what in an earlier 
chapter was called the promorphology of the egg are of this 
sort. There is one kind of promorphology that is of special 
importance to the present stage of this discussion. I refer 
to the kind known sometimes as "germinal localization" and 
sometimes as "organ forming substances" in the ovum. The 
idea, expressed in a sentence, is that in the eggs of some 
animals, portions of the egg destined to give rise to par- 
ticular parts of the future embryo are visibly different from 
other portions before cell division begins, in some cases even 
before maturation and fertilization occur. According to 
our understanding of heredity, these distinguishable por- 
tions of such eggs are themselves hereditary attributes not 
only of the animal species to which the eggs belong, but of 
the eggs, no less than are distinctive morphological features 
of the adult animals or of any developmental stages. The 
study of these attributes of eggs is peculiarly interesting 
since, belonging to germ-cells par excellence, if we can get 
observational evidence on both their origin and destination, 
we shall have direct evidence that one and the same substance 
is determined on the one hand by heredity, and on the other 
is a determiner in the strict genetic sense of hereditary at- 
tributes yet to be developed. 

(a) Eggs of Ascidians--The Facts 

Because of the great importance of the observations of 
E. G. Conklin in this field, and of his general views con- 
cerning the bearing of his observations on heredity, we shall 
make his work the center of our examination. One of the 
most important of Conklin's investigations was on the eggs 

Evidence from Metazoan Germ-Cells 


of several species of Ascidians. Among the great merits of 
this investigation are the facts that the normal living eggs 
were studied with great care, and that the comparative 
method was employed to a considerable extent. 




C rt. 



cn., chorion. c.p., clear protoplasm, c.r., crescent of mesoderrnal 

substance, g.v., germinal vesicle, p.b., polar body, p.l., peripheral 

layer of protoplasm, t.c., test cells, y.h., yellow hemisphere of 
egg- yk., yolk. 

Three species of simple Ascidians, Ciona intestinalis, Mol- 
gula manliattensis and Styela partita furnished the eggs 
for the investigations. 

We will begin the examination by quoting from the sum- 

18 The Unity of the Organism 

mary of results, and then use these as the basis of a closer 
scrutiny of the conclusions. Under the head "Organization 
of the Egg" we find : "14. In the ovocyte of Cynthia partita 
there is a peripheral layer of yellow protoplasm, (p. I.) a 
central mass of gray yolk, (//A 1 .) and a large clear germinal 
vesicle (g. v. ) which is eccentric toward the animal pole 
(figure 43a). These same parts are present in the eggs 
of other ascidians, but are differently colored. 

"15. When the wall of the germinal vesicle dissolves at 
the beginning of maturation divisions a large amount of 
clear protoplasm, containing dissolved oxychromatin, is 
liberated into the cell body. This clear protoplasm is ec- 
centric toward the animal pole and is distinct from the yolk 
and peripheral layer. 

"16. Immediately after the entrance of the spermatozoon 
the yellow and clear protoplasm flow rapidly to the lower 
pole, where the yellow protoplasm collects around the point 
of entrance ; the clear protoplasm lies at a deeper level. The 
yellow protoplasm then spreads out until it covers the sur- 
face of the lower hemisphere. This flowing of protoplasm 
to the point of entrance of the sperm is comparable with 
what takes place in many animals, though much more ex- 
tensive and rapid here than elsewhere (figure 43b). 

"18. The sperm nucleus moves from the point of en- 
trance toward the equator in a path which is apparently 
predetermined. This path lies in the plane of first cleavage 
and the point, just below the equator, at which the sperm 
nucleus stops in its upward movement, becomes the posterior 
pole of the embryo. The median plane and the posterior 
pole are probably not determined by the path of the sperma- 
tozoon but by the structure of the egg. All the axes of the 
future animal are now clearly established antero-posterior, 
right-left, dorso-ventral. 

19. The yellow protoplasm "collects into a yellow cres- 
cent with its middle at the posterior pole and its horns ex- 

Evidence from Metazoan Germ-Cells 19 

tending about half way around the egg just below the 
equator (figure 43c. yji.) This position it retains through- 
out the whole development, giving rise to the muscle and 
mesenchyme cells (figure 43 d. cr.) 

"20. . . . At the close of the first cleavage (figure 43d) 
the nuclei and clear protoplasm move (c.p.) into the upper 
hemisphere and thereafter, throughout development, this 
hemisphere contains most of the clear protoplasm and gives 
rise to the ectoderm. 

"21. . . . When the clear protoplasm moves into the 
upper hemisphere the yolk is largely collected in the lower 
hemisphere. This yolk-rich area gives rise to the endoderm. 

"24. The chief factor of localization is protoplasmic 
flowing ; cell division is a factor of subordinate value." 7 

From the numbers given in this quotation it will be recog- 
nized that only a portion of the summing up of results is 
here presented; and it will be understood that each item in 
the summary represents a lengthy, detailed description in 
the body of the memoir. This summary we may again sum- 
marize as follows : By the time the first division of the egg- 
cell is completed the portions of the egg which are to give 
rise to three great groups of tissues of the future animal 
are distinguishable from one another by definitely visible 
attributes. The clear protoplasm situated at the upper 
pole of the cell will give rise to the external epithelium or 
skin; the yolk-laden protoplasm in the lower hemisphere will 
produce the epithelial lining of the digestive tract ; and 
from the yellow protoplasm gathered on the surface at the 
lower pole will come most of the muscle and connective tis- 
sue of the animal. 

(b) Conklin's Interpretation 

Passing now to a consideration of these facts in relation 
to heredity we must not neglect to notice the two-fold aspect 

20 The Unity of the Organism 

of the question, namely that we have to do on the one hand 
with attributes of the egg itself which are results or termini 
of the kind that gave rise to the conception of heredity ; 
and on the other hand with these same attributes not as 
results but as causes or 'forerunners of later-appearing at- 
tributes, also by the same criteria due to heredity. That the 
egg attributes with which we are dealing are due to heredity 
is obvious from the fortunate circumstance that, as already 
indicated, the observations under review were comparative 
to considerable extent. Thus we have this general com- 
parison of the unripe eggs of the three species : "In the liv- 
ing eggs of Cynthia this peripheral layer is clear and trans- 
parent and contains uniformly but sparsely distributed 
yellow pigment, which seems to be associated with these small 
refractive spherules. ... In Ciona and Molgula also these 
three areas are distinguishable in the living egg before ma- 
turation, but not so clearly as in Cynthia. In Ciona the 
peripheral layer is nearly transparent, the yolk is a brown- 
ish red. ... In Molgula both the peripheral layer and the 
germinal vesicle are transparent, while the yolk is gray, 
with a faint lilac tinge." 8 As an example of a very definite 
statement of specific difference between the eggs we read, 
"In Ciona the same type of protoplasmic movement occurs 
as in Cynthia, but with certain minor differences." 

The evidence brought forward by Conklin of species differ- 
ences in eggs is not restricted by any means to Ascidians. 
In some of the gastropod mollusca, for example, similar re- 
sults were reached by a very notable research on the quanti- 
tative relations of various egg organs of several species of 
the genus Crepidula. To be explicit in a single case only, 
the relative volume of macromeres 3A-3D to 3a-3d was 
found to be 12.1:1 in Crepidula plana and 59.3:1 in C. 
convex. From a long series of determinations of difference, 
Conklin remarks : "One cannot study the eggs of different 
animals without being much impressed with the fact that 

Evidence from Metazoan Germ-Cells 

the distribution of yolk to the four macromeres is highly 
characteristic of different species and orders." Yet, as 
we shall have occasion to point out later, it seems certain 
that facts of this kind have deeper meanings than Conklin 
has appreciated. 

After all that has been said in previous pages about 
specific differences, the point aimed at here will be obvious. 
The "decidedly thicker" layer of peripheral protoplasm at 
the lower pole in dona than in Styela, and the brownish 
red yolk in dona as contrasted with the white yolk tinged 
with lilac in Molgula, are indeed "minor differences." But 
species differences in adults and in all other stages are minor 
as a rule. The difference between the transparent-white 
color of the full-grown Ciona intestinalis and the pale, green- 
ish yellow of the adult Molgula manhattensis is also minor 
as contrasted with the "same type" of organization of the 
two animals. But it is just this being true to type as re- 
gards minor differences in living beings that has given rise 
to the conception not only of organic species, but to a large 
extent, of heredity as well ; and how can any one recognize 
the differential color of the grown-up Ciona as compared 
with the grown-up Molgula as due to heredity, but refuse 
to recognize the differential color of the eggs of the two 
animals as due to the same cause? 

We are, then, bound to accept the attributes of the eggs 
as the results of heredity, and so inquire about the "physi- 
cal basis" or "bearers" of these attributes. And this brings 
us again to the main issue of this section: Have we or have 
we not observational evidence that any other substance or 
substances than chromatin contribute to the production of 
the egg-phenomena under consideration? 

Conklin's conclusions and mode of reasoning as to the 
bearings of his observations on heredity are so crucial for 
the point now engaging us that we must examine them in 
considerable detail. Returning to the summary of results 

22 The Unity of the Organism 

upon which we have already drawn we read : "26. The 
organization of the ovocyte is not the initial organization. 
The yellow protoplasm (mesoplasm) of the Cynthia egg is 
probably derived, at least in part, from sphere material 
(archoplasm) which arose from the nucleus at the last 
ovogenic division. The yolk (endoplasm) is formed by the 
activity of the 'yolk matrix' (Crampton) which also is 
probably sphere material. The clear protoplasm (ecto- 
plasm) is derived from the germinal vesicle at the first 
maturation division. Thus many important regions of the 
egg come, at least in part, from the nucleus, and a method 
is thereby suggested of harmonizing the facts of cytoplasmic 
localization with the nuclear inheritance theory." 

Again, we find : "This truly remarkable condition in which 
considerable portions of the cytoplasm are traceable to the 
nucleus is of the utmost theoretical importance. From all 
sides the evidence has been accumulating that the chromo- 
somes are the seat of the inheritance material, until now this 
theory practically amounts to a demonstration. On the 
other hand, all students of the early history of the egg have 
observed that the earliest visible differentiations occur in 
the cytoplasm, and that the position, size and quality of 
the cleavage cells and of various organ bases are controlled 
by the cytoplasm. However, in the escape of large quanti- 
ties of nuclear material into the cell body, and the forma- 
tion there of specific protoplasmic substances we have a 
possible mechanism for the nuclear control of the cytoplasm, 
and when, as m the case of the ascidians and fresh water 
gastropods, these substances are definitely localized in the 
egg, and can be traced throughout the development until 
they enter into the formation of tlie particular portions of 
the embryo, a specific mechanism for the nuclear control of 
development is at hand, and the manner of harmonizing the 
facts of cytoplasmic organization with the nuclear inher- 
itance theory is clearly indicated." 

Evidence -from. Metazoan Germ-Cells 

(c) Critical Examination of Conklin's Interpretation 

One is struck at the outset of a critical examination of 
these sentences by the fact that what is claimed is that ob- 
servations have been made whereby a method is "suggested 
of harmonizing the facts of cytoplasmic localization with 
the nuclear inheritance theory" ; and that this suggested 
harmonization of sucli localization with the "nuclear in- 
heritance theory" (not "chromatinic inheritance theory" 
be it noticed) is the final count in a mass of evidence which 
"practically amounts to a demonstration" that the "chro- 
mosomes are the seat of the inheritance material." 

In other words objective facts which are only suggestive 
of a conclusion as touching the inheritance role of the whole 
nucleus rise to the demonstrational level as touching the 
same role of very small parts of the nucleus. 

The method by which this particular piece of logical 
sleight-of-hand is performed is easy to see, for though va- 
ried to meet the exigencies of the special case, it follows the 
general scheme of elementalistic interpretation with which 
we have become familiar. In the first place, the fact that 
the observations furnish very little if any direct evidence 
that the chromosomes cause the cytoplasmic flowing and 
localization is made innocuous as evidence against the chro- 
mosome dogma by assuming that the flowing and localization 
do not themselves come under heredity. That such an as- 
sumption is implied, in the argument seems certain from 
the fact that Conklin did not consider that the general 
theory of chromosomes as the bearers of heredity which he 
had espoused made it incumbent upon him to take cognizance 
of the fact that he himself had in reality testified that 
portions of the nucleus other than chromosomes are the 
seat of inheritance material. We have here another and a 
very notable case of shielding a prevalent theory by defini- 
tion ; that is, of shutting relevant but inimical facts away 

The Unity of the Organism 

from it by definition. That Conklin should have been un- 
wittingly led into this is the more surprising in that his 
own studies, particularly those on comparative morphology 
and physiology of the eggs of several groups of animals, 
have added largely to the proof that many of the gross 
attributes of the eggs themselves are subject to heredity. 
Indeed no biologist has expressed more positively than he 
the conception that the egg is the individual animal in one 
of its stages of development ". . . from its earliest to its 
latest stage an individual is one and the same organism ; 
the egg of a frog is a frog in an early stage of development 
and the characteristics of the adult frog develop out of the 
egg, but are not transmitted through it by some 'bearer of 
heredity.'" 12 

One wonders if Conklin really would maintain that only 
the attributes of an animal in the adult stage of its life are 
subject to heredity; or even that the attributes of any of 
its developmental stages are not so subject, were he con- 
fronted with the question in this form. Although I have 
been unable to find statements in his writings from which 
one may positively infer what his answer would be, yet 
several passages can be brought together which can not, I 
believe, be harmonized with one another, but reveal real 
contradiction. Thus in The Mechanism of Heredity, al- 
ready cited, we find : "Differentiation, and hence heredity, 
consists in the main in the appearance of unlike substances 
in protoplasm and their localization in definite regions or 
cells. Such a definition is as applicable to the latest stages 
of differentiation, such as the formation of muscle fibers, 
as it is to the earliest differentiations of the g-erm cells, 

o 7 

and the one is as truly a case of inheritance as is the other. 
In short, different substances appear at an earlier or later 
stage in the development of all animals, and these substances 
are then sorted out and localized; this is differentiation 
= heredity: see above]. Physiological division of labor in- 

Evidence from Metazoan Germ-Cells 25 

volves morphological division of substance; sorting out of 
functions implies sorting out of the material substratum 
of functions." 

If cytoplasmic sorting out and localization as well in the 
earliest as the latest stages of development constitute dif- 
ferentiation "and hence heredity," how, one must ask, can 
the hypothesis that "the chromosomes are the seat of the 
inheritance material" ever "practically amount to a demon- 
stration"? The only way, so far as I can see, to reconcile 
these two statements is to say that in so far as the expres- 
sion "seat of inheritance material" means anything, both 
chromosomes and cytoplasm are such seats, and hence that 
neither is the seat of it. 

But it is not enough to point out that there is contradic- 
tion here. We must try to discover just how so careful a 
reasoner as Conklin should fail to detect it, for we may 
feel certain that the failure is due not to mere oversight or 
carelessness but to some defectiveness in standpoint or gen- 
eral mode of procedure. Conklin fully realizes, as our quo- 
tations show, that movements of the cytoplasm go far to- 
ward determining the attributes of the eggs of Ascidians 
and many other animals. But the following makes the 
recognition still more definite: "Undoubtedly the most im- 
portant of all the localizing factors so far recognized are 
cytoplasmic movements." 14 

Assuming that our contention is valid that these localiz- 
ing factors of the cytoplasm are inheritance factors (and 
the virtual admission of this by Conklin in one of the two 
statements which we hold to be contradictory should be 
noted) we have still to see by what facts and reasoning Conk- 
lin reaches the view that his observations support the 
theory that "chromosomes are the seat of inheritance mate- 
rial." The observations in this case which support the 
chromosome theory are that the three kinds of cytoplasm 
of the egg: the yellow protoplasm (mesoplasm), the sphere 

The Unity of the Organism 

material (archoplasm), and the clear protoplasm (ecto- 
plasm) come, at least in part, and at one time or another, 
from the nucleus. It is not claimed that the chromosomes 
or even chromatin can be observed to produce the cytoplas- 
mic localization or any of the other distinctive features of 
the egg, as for example the different colors characteristic 
of different animal species ; or the different kinds of proto- 
plasm in the same species. Just what can be observed as 
to the relation between the chromosomes and the several 
kinds of material which pass out of the nucleus into the 
cytoplasm is not much dwelt upon by Conklin in this investi- 
gation. The clear protoplasm (ectoplasm) he describes and 
figures very definitely as being the major clear mass of the 
germinal vesicle set free in the cytoplasm as the nuclear 
membrane dissolves at the beginning of maturation. The 
chromosomes are said to be distinguishable at this time as 
numerous small deeply staining bodies. They can be ob- 
served to collect together in the "center of the nuclear 
area" and certain things can be made out about the shapes, 
and arrangement relative to one another of the individual 
chromosomes ; but nothing is recorded to indicate that they 
take any part in the movements of the ectoplasm, much less 
in the production of it. 

Concerning the relation of the other two kinds of cyto- 
plasm, the yellow or mesoplasm and the sphere material or 
archoplasm, to the chromosomes, Conklin's description is 
still more meager. An investigation by Crampton, however, 
has given particular attention to the relation of the sphere 
material, or as he calls it the yolk-matrix, to the chromatin. 
Crampton's results are the more significant for this discus- 
sion in that Conklin was undoubtedly acquainted with them. 
As the result of several searching tests of the chemical 
character of the yolk-matrix, Crampton says : "In Molgula, 
certainly, these granular masses are not chromatin in the 
proper sense." Although Crampton believes the substance 

Evidence from Metazoan Germ-Cells 

called by him yolk-matrix and considered by Conklin to be 
the same as what he calls sphere material, arises from the 
nucleus, this belief rests not on direct evidence of the pass- 
age of the substance from the nucleus into the cytoplasm, 
but on the facts that when the substance is first seen in 
the oocyte it is a small mass situated in the cell-body or 
cytoplasm close in contact with the nuclear wall, and that it 
reacts to stains and digestive fluids in the same way that 
certain granular contents of the nucleus react. But Cramp- 
ton is very explicit in pointing out that this substance in the 
nucleus is not chromatin, at least of the ordinary kind. 
Upon treating the cells with digestive fluids it disappears 
from both inside and outside the nucleus, the true chromatin 
being then left in clear view as fine granules in the nuclear 

So far as concerns the endoplasm, then, even though it 
be derived from sphere material and this in turn from the nu- 
cleus, the most trustworthy evidence we possess is to the 
effect that its primal nuclear source is not chromatin. In 
view of such facts as these, made known by Crampton and 
others, through what reasoning would Conklin, to whom the 
facts are familiar, still hold that taken all-in-all they sup- 
port the chromatin dogma of heredity? Readers whose 
minds have become sensitized to the general type of reason- 
ing which pervades nearly all elementalistic theorizing and 
makes it to some extent fallacious, will readily anticipate 
about how the argument will run in this case. But it will 
be profitable to see it in actuality. After saying that "some 
of the important cytoplasmic substances can be actually 
seen to come from the nucleus" but that "this does not indi- 
cate that these substances exist from the beginning in the 
nucleus ; on the contrary there is direct and visible evidence 
that they arise epigenetically," Conklin continues : "Such 
epigenesis, however, does not signify lack of primary organ- 
ization ; on the other hand all the evidence favors the view 

8 The Unity of the Organism 

that back of the organization of the cytoplasm is the or- 
ganization of the chromosomes, which is definite,, determinate 
and primary." (Italics mine). 

There you have it again ! Although it is freely granted 
that you can see the cytoplasm in the very act of arising 
epigenetically and moving about to become definitely located, 
that is, to become organized, still what you see is no part of 
the real essence of the business. "Back" of this, in the 
chromosomes, which, be it specially noticed, can not be seen 
to take any active part in the operations, we must conceive 
is the "organization" which is "definite, determinate and 
primary" -in other words which is The Ultimate Cause, so 
far as heredity is concerned. Again I repeat, wearisome 
as the iteration has become, that the fallacy in this sort 
of reasoning is not in holding that there is some causal 
power "back" of the phenomena to be explained, but that 
all such power is located there. That is, stating the general 
point in its application to the special case, the fallacy lies 
not in holding that the chromosomes contribute something 
to the hereditary attributes of the ascidians and other ani- 
mal groups whose development Conklin investigated, but in 
the implied denial that the cytoplasm contributes anything 
to it. 

Conklin probably would not admit that there is real 
contradiction between the observations by himself and oth- 
ers, on the part played by cytoplasm in the early stages of 
development, and his contention that the evidence now "prac- 
tically amounts to a demonstration" of the correctness of 
the theory that the "chromosomes are the seat of the in- 
heritance material." What he would probably contend is 
that the observations are opposed merely to the extreme 
form of the chromosome theory. Thus, speaking from an 
angle of the general subject a little different from that of 
cytoplasmic localization, he writes : "This conclusion is not 
a refutation of the nuclear inheritance theory, but it is a 

Evidence from Metazoan Germ-Cells 

profound modification of it." 17 And still more recently 
he has said, "Many biologists maintain that the nucleus and 
more particularly the chromosomes are the exclusive seat of 
the 'inheritance material' and that all the 'determiners' of 
adult characters are located in them. Against the extreme 
form of this theory many general and specific objections 
may be urged. General objections are based upon the con- 
sideration that the entire cell, cytoplasm as well as nu- 
cleus, is concerned in differentiation and that neither is 
capable of embryonic development in the absence of the 
other. Differentiation is indeed the result of the interaction 
of nucleus and cytoplasm, and how then can it be said that 
the nucleus is the only seat of the inheritance material?" 

An elaborate discussion of whether the language here used 
can be harmonized with the statement quoted above about 
the demonstration of the correctness of the chromosome 
theory, by saying that the views expressed in the last quota- 
tion merely involve a "profound modification of the nuclear 
inheritance theory" would smack too much of pure dialectics 
to deserve a place in this volume. Our sole concern is with 
the truth about the thing itself. 

Conklin's position would be so far satisfactory if he 
would permit us to understand his statements "the entire 
cell, cytoplasm as well as nucleus, is concerned in differentia- 
tion," and the one about modification of the nuclear inheri- 
ance theory, to mean that cytoplasm is "inheritance mate- 
rial" and contains "determiners." Evidence that cytoplasm 
contains "determiners" is even more positive than is that 
for the theory that chromosomes are the seat of such things, 
for the simple reason that we can observe abundantly cyto- 
plasm in the act of producing hereditary structures, whereas 
we rarely observe chromosomes operating directly in this 
way. But such permission would not, I fear, be forthcoming. 
If it would be, one is at a loss to understand why the terms 
"hereditary substance," "physical basis of heredity," "de- 

30 The Unity of the Organism 

terminers," "factors" and the like, constantly used in con- 
nection with the chromosomes are never used in connection 
with cytoplasm. Indeed, so well does Conklin present the 
general argument for the participation of cytoplasm in 
the development of hereditary structures that it is surpris- 
ing, not to say disappointing, to find him neglect to present 
the most specific argument we have to the same effect, name- 
ly that the genesis of a vast range of such structures can 
be directly observed to be largely due to cytoplasmic trans- 

One other passage in Conklin's general argument is so 
significant that we must reproduce it. "Differentiation is 
indeed the result of the interaction of nucleus and cyto- 
plasm, and how then can it be said that the nucleus is the 
only seat of the inheritance material? If held rigidly, this 
theory involves the assumption that the cytoplasm and all 
other parts of the cell are the products of the chromosomes, 
and that therefore the chromosome and not the cell is the 
ultimate independent unit of structure and function ; an 
assumption which is contrary to fact. Furthermore, since 
heredity includes a series of fundamental vital processes 
such as assimilation, growth, division and differentiation, 
there is something primitive and naive in the view that this 
most general process can be localized in one specific part of 
the cell, something which recalls the long-past doctrine that 
the life was located in the heart or in the blood, or the 
ancient attempts to find the seat of the soul in the pineal 
gland or in the ventricles of the brain." 19 

This passage contains several well-sent shafts not only 
against chromosomal elementalism, but against the elemen- 
talist standpoint generally. And I must recur again in con- 
nection with it, while the facts of egg organization as pre- 
sented by Conklin are fresh in mind, to the perception, 
indicated in previous chapters, that the physical-chemistry 
conception of the cell must be extended to the organism. If 

Evidence from Metazoan Germ-Cells 31 

Conklin once sees the full force of this contention, he will, 
we may hope, be ready to let go entirely of the idea that 
the facts of cytoplasmic organization must be "harmonized 
with the nuclear inheritance theory." He will then see 
that there is no more necessity for harmonizing the facts of 
cytoplasmic organization with the nuclear inheritance the- 
ory than there is for harmonizing the facts of nuclear or- 
ganization with the theory of cytoplasmic inheritance. 


1. Ballowitz 255 11. Conklin ('05) 101 

2. Wilson, E. B. ('00) 135 12. Conklin ('08) 90 

3. Ballowitz 277 13. Conklin ('08) 92 

4. Oliver 489 14. Conklin ('05) 102 

5. Oliver 479 15. Crampton ('99) 48 

6. Casteel 646 16. Conklin ('05) 101 

7. Conklin ('05) Ill 17. Conklin ('08) 98 

8. Conklin ('05) 11 18. Conklin ('15) 162 

9. Conklin ('05) 21 19. Conklin ('15) 163 

10. Conklin ('05) 114 

Chapter XV 



WE must now give the greatest possible concreteness 
to the general truths stated by Conklin that growth 
is the result of the interaction between nucleus and cyto- 
plasm and that heredity includes such fundamental vital 
processes as assimilation, growth, division, and differentia- 
tion. But the one and only way, I again insist, to attain 
concreteness and certainty in the matter is through a maxi- 
mum of observation coupled with a minimum of inference. 
That is, the goal must be reached mainly by direct study of 
the anatomical, histological and physiological transforma- 
tions through which hereditary attributes are produced. The 
issue can be met squarely only by a still further considera- 
tion of what we actually know about the participation of 
all sorts of elements of relatively undifferentiated cells in 
the production of obviously hereditary parts. The study 
of interactions between nucleus and cytoplasm and of as- 
similation growth, etc., in germ-cells is not enough. What 
we have seen in preceding pages about the development of 
organs in the protozoa and in spermatozoa is that much 
toward the end sought. Our task now is to consider the 
local transformations by which structures are produced in 
multicellular organisms, especially in those which develop 
from eggs. 

The Mitochondria! Theory of Heredity 

This task may well begin by examining the recent efforts 
to locate the "hereditary units" in the mitochondria of the 


Evidence from Somatic Hist o genesis 33 

cytoplasm instead of in the chromosomes. This effort is 
just as misdirected as is the effort to make chromatin the 
sole hereditary substance. According to the organismal 
conception, all life phenomena, including those of inheri- 
tance, consist in the activities and interactivities of an 
enormous number of substances and units and forces, all of 
which, in exhaustive analysis, are dependent upon the organ- 
ism as a living whole. It is, therefore, as futile to hunt in 
one corner as another for the physical basis of heredity 
in an exclusive and more or less metaphysical sense. Any 
one who has grasped this idea will know beforehand that 
the proposal to make mitochondria fully explain heredity is 
doomed to failure no less certainly than was the proposal to 
make chromosomes or any one kind of cell element play such 
a role. But hypotheses, the falsity of which might have been 
seen before they were tested, may still be useful. If those 
who propose them can be convinced of their fallacy in no 
other way than by testing them then it is better that they 
should be tested even though much time and labor be given 
to the task. Again, the evidence brought out which dis- 
proves an hypothesis may be highly useful in establishing 
some alternative general view. This result has been espe- 
cially striking in the case of the mitochondrial hypothesis of 
hereditary substance. By bringing the cell-body back into 
the field of interest, from which it had been largely excluded 
so far as heredity was concerned, by the nuclear inheritance 
theory, the mitochondrial hypothesis has resulted not merely 
in proving that mitochondria are not bearers of heredity 
in the elementalist sense, but that in a rational sense they 
are sometimes rather closely concerned in the production 
of hereditary structures ; and, of even greater importance, 
that still other cytoplasmic material is likewise so concerned. 
The only reason why the mitochondrial theory of hered- 
ity is less interesting than the chromatin theory is that 
there is so much less observational evidence in support of it. 

34 The Unity of the Organism 

In fundamental principle the one is no more and no less 
acceptable than the other. 

Any half plausible suggestion that a particular minute, 
obscure part of the germ-cell may be a "bearer of heredity" 
seems to endow that part with a peculiar fascination to biol- 
ogists who have the elementalist habit of thinking, and this 
secures to it an inordinate attention until the untenability 
of the hypothesis is so overwhelmingly proved that even the 
most credulous are forced to abandon it. 

Several biologists have recognized something of the state 
of mind here indicated without, however, perceiving its real 
meaning. Thus in a review of the work of Meves, which 
will be examined presently, we read: "the new interpretation 
which Meves gives at this time indicates that many are 
still dissatisfied with the all-sufficiency of the [nuclear] 
theory, and are eagerly seeking and grasping, as it were, 
the first visible sign of any other substance which may serve 
to carry the hereditary qualities." The remark to be 
made about any statement of this kind is that the real 
though usually unperceived ground of dissatisfaction is not 
with the all-sufficiency of the nuclear theory of heredity, but 
with the all-sufficiency of any theory that attempts to local- 
ize the function of carrying heredity in some small, specific 
fraction of the germ-cells ; and that the attitude which 
Doctor Payne well characterizes as "eagerly seeking and 
grasping" which has marked so much of recent search after 
the physical basis of heredity, has a large measure of genu- 
ine illusion in it. Inspired by the ages-old, alluring belief 
that an imperceptible final cause and explanation is hidden 
somewhere within or behind whatever is grossly sensed, the 
pursuit becomes "eager and grasping," which is another 
way of saying that it becomes more or less irrational and 

The truth of these remarks is rather strikingly exempli- 
fied in the short, somewhat feverish history of the mitochon- 

Evidence from Somatic His to gene sis 35 

drial theory of heredity. The meager observational support 
for the hypothesis that these particular cell members are 
generally bearers of heredity relieves us from the necessity 
of examining it in any such detail as we examined the chro- 
mosome theory. But there are several things about it of 
so much importance that we must look into it somewhat. 
The name mitochondria was first used by Benda for a "new 
cell organ, perhaps serving a specific function." Benda's 
original view was that the function served is that of the 
motility of the cell. But in a later publication he presented 
observations which seemed to him conclusive proof that the 
mitochondria of the sperm are situated in parts of it which 
enter the egg at fertilization. This last suggestion gave 
an added impulse to the study of the bodies occurring in the 
cytoplasm and soon many new names were applied to them, 
for they were soon found to present differences in size, 
shape, and reaction to chemicals. It seems, however, that 
the present state of knowledge justifies us in applying to 
them all the one term, mitochondria, though without imply- 
ing that they are all exactly alike. They may be held to 
be generically alike but specifically different. 

The first investigator to set up definitely the hypothesis 
that cytoplasmic elements of this class are bearers of hered- 
itary qualities seems to have been Meves. 

Only a few of the very many investigations since devoted 
to the subject can be noticed, these being selected for their 
bearing on particular aspects of the problem. In the first 
place, the position of Meves himself is important. Accept- 
ing the assumptions formulated by O. Hertwig in 1875 of a 
"substance which carries over the beginnings (Anlagen) of 
the parents to the child," and that "this substance exists 
(in the germ-cells) in an original, histologically undifferen- 
tiated condition" ; 3 and adding his own reflection that not 
all the cytoplasmic parts of the spermatozoon (for example, 
the axial fiber of the tail) possess inheritance potencies, he 

36 The Unity of the Organism 

advanced the hypothesis that the mitochondria answer the 
conditions of Hertwig's theory for the cytoplasmic part of 
the male, though probably not of the female germ-cells. 
Thus Meves reached the rather attractive conception that 
"heredity is accomplished through protoplasm and nucleus 
together." Were we to know no more than this about his 
theory we might suppose his position to be that of a genuine 
integrationist. However, his very next sentence does not 
permit us to question any further the orthodoxy of his ele- 
mentalism. "The qualities of the nucleus," he says, "are 
carried over by the chromosomes, those of the plasma by 
the chondriosomes." In other words the working together 
of nucleus and cytoplasm which he conceives is not of the 
sort which makes the part played by each contribute to all 
the results, but that one set of elements produces its partic- 
ular part of the total effects, while another set produces 
another part of the effects. 

The Mitochondria} Theory Tested by the Ontogeny of the 


The utter inadequacy of this hypothesis is shown by some 
of the same evidence which revealed to us the inadequacy of 
the chromatin theory, namely that when we come to study 
the histogenetic processes by which innumerable hereditary 
attributes are produced, we find portions of the cytoplasm 
other than the mitochondria taking the leading part in the 
production. Perhaps no single one of the many instances 
examined by us of cytoplasmic participation in the produc- 
tion of attributes refutes Meves' hypothesis more complete- 
ly and instructively than that of the developing sperm of the 
fowl tick already described. 

We have already examined this case as one of special weight 
in proving that the cell-body ^ in contradistinction to the nucleus, 
is hereditary substance. Now we must see the conclusiveness 

Evidence from Somatic Hist ogene sis 37 

with which this particular ontogeny disposes of the mitochondria! 
hypothesis of heredity so far as this case is concerned. 

Recurrence to the description and figures (42 a, b, c, d, e, f ) will 
recall that the mitochondria, widely distributed through the cyto- 
plasm in the early spermatocyte (?HZ.) assemble into a rather 
sharply defined ring-shaped mass (m.r., figure 42b) as the cell 
transformation proceeds., and finally take a position in the devel- 
oping sperm about as remote as they can get from some of the 
most actively and fundamentally changing parts of the organism. 
(mi. f figures 42 e, f, g). 

There is no more possibility of explaining the development of 
many of the parts of this sperm, the outer sheath, for example 
(o.t., figures S6e and f ) as due to the influence of the mitochondria 
than as due to the influence of the nucleus. And I point out 
again for the hundredth time what the real issue is here. The 
observations certainly do not exclude the possibility that the 
mitochondria exercise some invisible influence on the development 
of, say, the outer sheath. There may be or there may not be such 
an influence. But the observations do show conclusively that 
cytoplasmic portions of the cell other than mitochondria are 
operative in producing the outer sheath. 

It should be said in concluding this reference to the sperm of 
the chicken tick that Casteel finds the mitochondria located finally 
in the end of the sperm opposite that which contains the nucleus ; 
and that this end goes ahead in locomotion, the motion being pro- 
duced by a circlet of mobile processes at this end. From this he 
believes that the contractile elements are mitochondrial in origin. 
The conjecture that the mitochondria of the spermatocyte take 
part in producing the motor elements of the sperm tail is perhaps 
strengthened by the observations of other students, notably Lewis 
and Robertson. These investigators were able to follow the mito- 
chondria in the ontogeny of the living sperm directly into the 
tail, where they transform into two equal threads situated along- 
side the axial filament. 

The Mitocliondrial Theory Tested by Histogenesis 

If this hypothesis that mitochondria in developing sperm 
cells give rise to the motion-producing structures of the 
sperm tail, then the mitochondria would be genuine "inheri- 
tance material" for these particular elements, the hereditari- 

38 The Unity of tlie Organism 

ness of the motile elements being especially striking in the 
case of the chicken tick sperm from the fact that the mode 
of locomotion in this spermatozoon is almost unique. But 
while the case of mitochondria and other non-nuclear parts 
of the cell in the development of the spermatozoa, ought to 
be conclusive that although mitochondria can not be "hered- 
itary units" in any general sense, they, as well as other 
cytoplasmic parts, may contribute to the production of 
hereditary structures, yet it would not be so accepted, prob- 
ably, by the most exacting theorists because such biologists 
would not allow that a spermatozoon, being unicellular, can 
have organs and parts which are subject to heredity "in 
the strict sense" (i.e. in the sense of the definition of hered- 
ity set up by these persons). We must, consequently, pro- 
ceed to the specific task of this section ; namely that of 
considering what is known about the part played by mito- 
chondria in the histogenesis of hereditary structures in 
multi-cellular organisms. 

Nearly all the studies centered upon the question of 
whether the mitochondria are bearers of heredity have gone 
on the assumption, quite inadequate according to my view, 
that the problem is to be solved by ascertaining whether or 
not the bodies are persistent cell organs, take a definite part 
in fertilization, and are contributed in equal quantity by 
the female and male germ-cells. In other words the assump- 
tion has been that the same criteria which have been relied 
upon to prove that chromosomes are bearers of heredity, 
must also be applied for deciding whether or not mitochon- 
dria have the same office. But numerous studies have also 
aimed to follow the mitochondria in the genesis of tissues, 
and herein lies the chief importance of investigations in 
this domain. Not only have they greatly increased our 
knowledge about the role played by various parts of the 
cell in histogenesis, notably of the cytoplasmic parts, but 
they have put us in possession of much precise information 

Evidence from Somatic His to genesis 39 

as to what "hereditary substance" is. 

The results and conclusions thus far reached are on the 
whole so diverse, often so conflicting, that any attempt at 
a general review of them would be useless for this discussion. 
However, certain of the results, actual or claimed, are im- 
portant. For example, at one extreme it is contended that 
the mitochondria are the immediate precursors of the most 
distinctive elements of ah 1 classes of adult tissues. Thus 
Meves : "All these differentiations (of embryonic cells into 
tissue cells) however heterogeneous they may be, arise 
through the metamorphosis of one and the same elementary 
constituent of the plasma, the chondriosomes. The chondri- 
osomes are the material substratum lying at the basis of the 
processes of differentiation, which become the specific sub- 
stances of the different tissues." As an extension of this 
view we learn from Lewis and Lewis and other reviewers 
that the following tissues are reported on the authority of 
a long list of workers to be produced by the mitochondria: 
fibrillae, myofibrillae, fibrillae of epithelial cells, corneous sub- 
stance, secretory granules, fat, leuco-, chloro- and chromo- 
plasts, the test substance in foraminifera, and various other 
tissue elements. 

But several investigators, notably E. V. Cowdry, have 
shown the inconclusiveness of the evidence on which the con- 
tention is based that neurofibrils originate from mitochon- 
dria. "There is no evidence," Cowdry says, "that mito- 
chondria are transformed into neurofibrils. . . . The mi- 
tochondria do not show, either by a variation in their 
morphology, staining reactions, or in any other fashion, 
. . . indications of being transformed into material of dif- 
ferent chemical composition." 7 Furthermore, lie shows that 
the mitochondria do not diminish in quantity in any way 
commensurate with the increase of neurofibrils, as the neu- 
roblasts transform into ganglionic cells. Eminently worthy 
of note, as bearing on our contention made some pages 

40 The Unity of the Organism 

back, that cytoplasm itself is hereditary substance, is Cow- 
dry's detailed description of neurofibrils as a "differentia- 
tion of the ground substance" of the neuroblasts. And in 
a later paper the same author makes a strong case of the 
view that while mitochondria are "associated in some way 
with the formation of many substances," it is highly im- 
probable that they transform into them. 

On the whole the tendency of the latest investigations 
appears to be to deny that the bodies produce, in a strict 
sense, any tissue elements. Thus as a result of their quite 
remarkable studies on mitochondria of living cells Lewis and 
Lewis say, following the enumeration given above : "The 
above theories seem impossible to correlate. It seems evi- 
dent that the mitochondria are too universal in all kinds 
of cells to have the function of forming any one of the 
above structures of differentiated tissue, and in the light of 
what cytological chemistry is known, it appears practically 
impossible for the mitochondria to form all the cell struc- 
tures mentioned above. In view of the fact that the mito- 
chondria are found not only in almost all animal cells but 
in plant cells as well it seems more probable that they play 
a role in the more general physiology of the cell." The 
idea that the mitochondria are primarily concerned with the 
metabolism of the cell appears to be gaining ground under 
the present comprehensive and critical methods of investi- 
gation that are being applied to them. 

The Untenable Hypothesis that the Cytoplasm of the Ovum 
is Inheritance Material for General but not for 

Special Characters 

A number of biologists have recently put forward the 
hypothesis that while the cytoplasm of the egg-cell may be 
"hereditary material" for certain of the general attributes 
of the organisms, chromosomes ' "carry" the hereditary, 

Evidence from Somatic Hist ogene sis 41 

of the more specific attributes. This conception has arisen 
from a considerable range of observations to the effect that 
for quite a time in the "early ontogeny of many animals some 
of the attributes of the embryo can be seen to come directly 
from the cytoplasm of the egg. Thus both Driesch and 
Loeb have taken special notice of the fact that, as expressed 
by Loeb, "when the protoplasm of the egg possesses a strik- 
ing pigment the larva will possess the same for some time 
at least" ; and that "if such an egg is hybridized with the 
sperm of a form whose egg is unpigmented, the larva will, 
of course, possess a 'maternal' quality which is due solely 
to the protoplasm (Driesch)". 1 * And in the same connec- 
tion, Loeb continues : "It is obvious, then, that during the 
first stages of development an influence of the protoplasm 
upon heredity may make itself felt, which will disappear as 
soon as the protoplasm of the egg has been transformed into 
the tissues of the embryo." One of the cardinal questions we 
have to consider may be formulated in connection with this 
last quotation: Have we a right to assume that because 
an obvious influence of the protoplasm upon heredity dis- 
appears on the transformation of the protoplasm into tis- 
sue, therefore all such influence of the protoplasm ceases? 
To answer this question through observations upon the 
protoplasm of the cells concerned just before, during, and 
just after the transformations is exactly the central aim 
of this section. I can not refrain from making use of 
another sentence from Loeb to aid in defining the problem 
more clearly. "It does not seem to me," he writes, "that a 
discussion as to the relative influence of protoplasm and 
nucleus upon heredity will prove very fertile, but that it is 
necessary to transfer this problem as soon as possible from 
the field of histology to that of chemistry or physical chem- 
istry." I quite agree that "discussion as to the relative 
influence of protoplasm and nucleus upon heredity" can not 
be very fruitful. But the grounds of my skepticism are 

The Unity of the Organism 

widely different from those of Loeb. According to my view, 
the question is not one to be settled by discussion at all, 
but by observation coupled with a measure of consistent 
reasoning. Assuming that I am right, to "transfer" the 
problem "from the field of histology" if this really means, 
as it seems to, that the problem should be taken away from 
histology, no matter whether to the field of chemistry or 
any other, would be to remove it all the further from ob- 
servation and plunge it so much the deeper into discussion. 
I have not the slightest doubt that chemistry, especially 
biochemistry pursued on the principles of physical chem- 
istry, will have to be made large use of before the fullest 
possible understanding of the mechanism of heredity is 
reached. But this use will have to go hand in hand not 
only with morphological studies on germ-cells, but as well 
on hosts of cells during the whole ontogeny. Chemical in- 
vestigation will have to supplement, it cannot supplant, it 
cannot even lead, histogenic investigation. If there is one 
thing made more positive than any other about heredity by 
modern study of the subject, it is that heredity is some- 
thing which pertains to the smaller taxonomic grades of 
organisms, races, varieties, species and so forth. It w r ould 
seem, accordingly, that hardly any suggestion for the study 
of heredity could be wider of the mark than one to trans- 
fer it from the only field which makes any pretense of in- 
vestigating the details of development, and taking it into 
a field like that of physico-chemical activity, which is no- 
toriously devoid of the very attributes without which there 
would be no such thing as heredity. Having once ascer- 
tained by observation as much as possible about how hered- 
itary attributes are actually produced, it will then be in 
order to learn as much as possible about the chemistry of 
the processes. Chemistry can do its share in solving the 
problems of heredity after and not before histogenesis has 
done its share. 

Evidence from Somatic Histogenesis 43 

Conklin has expressed more definitely than any other biol- 
ogist with whose writings I am acquainted, the idea men- 
tioned above, that cytoplasm "influences" heredity in early 
ontogenetic stages, and also influences adult attributes of 
the major taxonomic groups, but becomes inoperative in 
the later stages of development, the heredity of these being 
transferred to the nucleus. He says, "In short, the egg 
cytoplasm fixes the general type of development and the 
sperm and egg nuclei supply only the details." And fur- 
ther: "We are vertebrates because our mothers were 
vertebrates and produced eggs of the vertebrate pattern ; 
but the color of our skin and hair and eyes, our sex, stature, 
and mental peculiarities were determined by the sperm as 
well as by the egg from which we came. There is evidence 
that the chromosomes of the egg and sperm are the seat of 
the differential factors or determiners for Mendelian char- 
acters, while the general polarity, S} T mmetry, and pattern 
of the embryo are determined by the cytoplasm of the 
egg." If two points in this last quotation be viewed in 
the light of a large mass of relevant evidence not usually 
taken into account in recent discussions on heredity, and 
if strict consistency in the use of terms be maintained, the 
general conclusion will be quite different from that stated 
by Conklin. These two points, the conception of "differen- 
tial" and of "determiner," must now receive attention ; but 
first I will illustrate my position by a case presenting the 
kind of evidence to which I have referred. 

Species Attributes in Single Cells of Adult Organisms 

In general, this evidence comes from the field of histology, 
or more strictly histogenesis. The most convincing, because 
the most direct, evidence from this source is that pertaining 
to the development of hereditary structures in adult meta- 
zoa and metaphyta. The structures in such organisms 

The Unity of the Organism 

which are the most indubitably hereditary are those which 
distinguish the smaller but yet definite taxonomic groups. 
A little consideration will convince one that about the most 
crucial cases would be those in which the development of 
differential attributes could be traced directly to cell struc- 
ture and development. It so happens that vast as is our 
knowledge of histogenesis, the part of it which answers 
directly to the requirements here laid down is by no means 

The Spmules of the Ascidian Genus Styela 

The best instances I have been able to find are super- 
ficial appendages in some animals and plants, so small that 
they consist of a few cells or even of a single cell. One strik- 
ing instance of this sort has come to light in my own studies. 



n., nucleus. 

It is that of the minute spines which cover the inner surface 
of the branchial siphon in some species of the ascidian group 
of Styalids. Huntsman first called attention to the fact 
that each spinule is a single cell, and that at least in some 
cases the structures furnish differentiating attributes for 
species. Miss Forsyth and I have reexamined the point for 
Styela montereyensis and S. yakutatensis, and are able to 

Evidence from Somatic Histogenesis 


confirm Huntsman's results. As Huntsman had opportun- 
ity to study the matter in a larger number of species than 
Miss Forsyth and I have had, the following description is 
taken largely from his paper. Figures 44a, b, 45, 46 are 
from Huntsman and figure 47 is from Hitter and Forsyth. 
By comparing figures of what may be called the dorsal view 
(figures 44a, 45, 46) with the side view (figure 44b) it is 
seen that the distinctive feature about the cell which con- 
stitutes the spinule is a shield-like plate on one side of the 
somewhat elongated cell, the distal end of which projects 
more or less beyond the cell body, the whole resembling to 


some extent the end of a finger with its nail. The shield 
is harder than the rest of the ceU, and is probably com- 
posed of the same material as the general "test" of Ascid- 
ians, animal cellulose. The spinules are so placed that the 
basal portion is embedded in the surface layer of the test 
on the inside of the siphon, the shield being on the free side 
of the cell with its free edge pointed toward the lumen of 
the siphon. The specific attributes furnished by the 
spinules depend upon the shape and structure of the shields. 
The free edges may be truncate (figure 44, S. yakuta- 
tensis), or long-pointed (figure 46, S. greeleyi), or low- 
conical (figures 45, 47, S. plicata, S. montereyensis). 
Again the edge may be smooth (figures 46, 47, S. greeleyi 

The Unity of the Organism 

and A^. montereyensis) 9 or it may be serrate (figures 44, 45 
S. yakutatensis, S. plicata). 

Viewing this case in the light of considerations put for- 
ward on the preceding pages, the pertinent queries about 
the heredity of the shields almost ask themselves : What is 


the "inheritance material" that causes the shield to be short 
and truncate in S. yakutatensis and long-pointed in S. 
greeleyi; or that explains the serrated edge in S. yakuta- 
tensis and S. plicata as contrasted with the smooth edge in 
S. montereyensis and S. greeleyi? Is the "seat" of that ma- 
terial in the cytoplasm or the nucleus of the shield-produc- 
ing cell? Unfortunately we have no direct observational 
information about the genesis of the spinules. But the 

Evidence from Somatic Histogenesis 

indirect evidence which bears on the point favors over- 
whelmingly the view that the cytoplasm is chiefly responsible 
for the shield. Huntsman supposes the spinules to be de- 
rived from the cells of the cellulose tunic characteristic of 
ascidians. He may be right in this ; but it may be, too, that 
they are derived from the epithelial lining of the siphon. 
The matrix of the cellulose tunic is undoubtedly largely if 
not wholly produced by the ectodermal cells. It is usually 
held to be secreted by these cells; but in some cases, in 



Perophora for example, a portion of the cytoplasm of the 
cells seems to become transformed into the cellulose. The 
process of transformation can be particularly well seen in 
the cells which line the branchial siphon of developing blas- 
tozooids as shown in figure 32, plate III of my memoir. 12 
The parts of the cell-bodies turned toward the cellulose are 
here long drawn out and the protoplasm gradually becomes 
indistinguishable from the surrounding cellulose substance 
in which it is imbedded. If now we imagine this protoplasm 
to transform not into the characteristic cartilage-like cellu- 
lose mass spread over nearly the whole surface of the body, 
that from innumerable cells fusing into a common mass, but 
each cell to retain its individuality, its protoplasm becoming 
the shield of a spinule, we should have what these styelids 

48 The Unity of the Organism, 

actually present. 

But whatever be the cells which transform into the spin- 
ules, all the available evidence indicates that the cytoplasm 
is the chief source of the shield part of the spinule. The 
possibility is not excluded that the nuclear chromatin also 
plays some part in the development. Indeed investigations, 
notably those by Duesberg on the ascidian egg made since 
"cytoplasmic inclusions" have come into prominence are 
distinctly favorable to such an hypothesis. But this is very 
far from proving that such chromatinic influence, assuming 
it to exist, reduces the cytoplasm to pure passivity. 

In the light of what is here set forth let us examine the 
view expressed by Conklin and quoted above that the egg 
cytoplasm fixes the general type of development, while the 
nuclei of egg and sperm together "supply only the details." 
The examination should be the more cogent from the fact 
that Conklin's idea was based largely on his investigation 
of ascidian eggs, some of which pertained to the very same 
genus, Styela, as do the spinules just described. The reader 
should not fail to notice that both Conklin and myself are 
dealing with single cells, he having to do with egg-cells, while 
I am concerned with spinule cells. Nor should the fact be 
lost sight of that these two categories of cells stand at 
about opposite extremes in the life career of an individual 
Styela, the egg being at the very beginning, while the spinule 
is produced at or near the completion of adulthood. Since, 
however, both cells possess attributes distinctive of the 
species, there can be no more "filling in of details" at one 
end of the series than at the other, so far as concerns the 
differential attributes under consideration. 

Now comes the main point. The differential attributes 
of the egg-cell and early embryo, their "polarity, symmetry, 
and pattern," are "determined by the cytoplasm," to use 
Conklin's own words. What is the evidence that these attri- 
butes are thus determined? That of direct observation, 

Evidence from Somatic Histogenesis 49 

as the examination of Conklin's work has shown. But if on 
the basis of such observation it can be asserted that attri- 
butes of egg and early embryo are determined by the cyto- 
plasm, how escape asserting that the same sort of evidence 
touching the production of the adult attributes, those per- 
taining to spinule cells, are likewise determined by the cyto- 

It was remarked above that if all known relevant facts 
were taken into account and consistency in terminology be 
maintained, Conklin's statement to the effect that we are 
vertebrates because our mothers were vertebrates and pro- 
duced eggs of the vertebrate type, but that our species and 
racial characters, color of skin and hair, and so forth, are 
determined by the chromatin of both parental germ-cells, 
would have to be greatly modified. We are now in position 
to see what modification is necessary. Although the state- 
ment is undoubtedly true that we are vertebrates because 
we develop from vertebrate eggs, the implication that the 
attributes which identify us with the human species and the 
Caucasian race are explained, so far as heredity is con- 
cerned, by the chromatin of the germ-cells, whether male or 
female or both, is not in accordance with all the observed 
facts bearing on the problem. The same kind of evidence 
on which the assertion is based that the embryonic charac- 
ters are determined by the egg cytoplasm, requires the 
assertion that skin and other adult characters are deter- 
mined by the same means. 

This leads to the remarks we have to make about con- 
sistency in the use of terms. The critical reader will hardly 
have failed to notice the difference in application of the 
word "determined" as used by Conklin in the quotation 
we are examining. When it is said that the color of skin, 
hair, stature and so on are determined by the germ-cells, 
the determining act or condition is far removed from that 
which is determined, and no direct causal connection be- 

50 The Unity of the Organism 

tween the two is established. On the contrary in the state- 
ment that the polarity, symmetry and pattern of the egg 
are determined by the cytoplasm, the determination is im- 
mediate and observed. Manifestly in a literal sense "de- 
termined" is properly used in the second connection but not 
in the first. The cytoplasm is operative on the spot, so to 
speak, in the second case. It is concerned with an immedi- 
ate result. The germ-cells on the other hand are not really 
determiners. They are not concerned with an end result, 
but are if anything instigators of a long developmental 
series at the far end of which appear the attributes in 
question: skin, color and the rest. 

If this case of spinule production stood alone as an 
instance of specific characters in adult animals traceable 
to cytoplasmic activity of individual cells, it would be a 
rather small base on which to erect a general argument in 
favor of cytoplasm as inheritance material. But it does 
not stand alone. Indeed, the company to which it belongs 
will almost certainly be found to be legion when systematic 
investigation of the subject shall have been made. 

The Spicules of Sponges and Other Invertebrates 

I will cite a few more cases. In several widely separated 
groups of animals, spicules, usually either calcareous or 
silicious, are present in some of the tissues. These are often 
produced by one or a very few cells, and often, too, their 
shape, size, and probably other attributes differ from 
species to species even of the same genus. 

The spicular system reaches its greatest development and 
has been most studied in sponges. "The spicules of 
sponges," writes Sedgwick, "in the diversity, symmetry, and 
intricacy of their form, in the perfection and finish of their 
architecture, constitute some of the most astonishing ob- 
jects in natural history." 13 Figure 48 gives a hint of the 

Evidence from Somatic Hist agenesis 


facts on which this statement is based. Although the prob- 
lem of what all these spicules are for does not directly con- 
cern us, indirectly it does, as the following further remarks 
of Sedgwick will indicate. "While it is pretty clear," he 
says, "that the main function of the skeletal structures is 
the support and protection of the sponge body, it is by no 
means easy to give explanations of the diversity and com- 


plexity of form which they present. The form of the megas- 
cleres is probably connected with the form of the canal sys- 
tem with which they are in relation (F. E. Schulze) ; but the 
form and even the existence of the microscleres defies any 
reasonable explanation." And then comes this statement, 
highly significant for almost any discussion of heredity : 
"By some spongologists the small spicules are regarded as 
functionless, and as having on that account a greater value 
for classificatory purposes." 

If any one wishes to be convinced of the extent to which 
the spicule forms differ with different species, he should 

The Unity of the Organism 

consult such systematic monographs as those of Schulze 
and Sollas in the reports of the Challenger Expedition. A 
picture of such a group of spicules as that shown in figure 
48 reminds one of pictures of ice crystals he has seen; and 
the question may well be raised, Are not these spicules in 
reality crystallization forms, and hence as devoid of hered- 
itary significance as are snow flakes? The fact that the 
form they have depends on the particular group of sponges 
to which they belong, i.e., that they follow the rules of 
biological taxonomy, is very strong evidence that they are 


genuine organic productions, and not mere crystallizations. 
And the further fact that they follow this rule even though 
many of them appear to have no functional significance in- 
dicates that their particular forms are due to heredity and 
not to modeling by extraneous influences in each individual 
sponge. But we are not left to such general evidence for 
support of the supposition that they are true organic 
products and subject to heredity. Their development has 
been studied by several zoologists and the results leave no 
doubt about their nature so far as this point is concerned. 

A single instance will be enough for our purpose, but it should 
be remarked that many others could be given. This is taken from 
the excellent summary of what is known about sponges written 
by Minchin for Lankester's Treatise on Zoology. 'To form a 
triradiate spicule three cells migrate into the parenchyma from 

Evidence from Somatic Hist ogene sis 

the dermal epithelium and become arranged in a trefoil-like figure 
(Figure 49). The nucleus of each cell then divides into two, in such 
a way that one nucleus is placed more deeply and one more super- 
ficially. Between each pair of sister nuclei a minute spicule ray 
appears, the three rays being at first distinct from each other but 

FIGURE 50 SEE 49. 

soon becoming united at the center of the system (Figure 50 tr. 
syst.}. As the rays grow in length the protoplasm of each actino- 
blast becomes aggregated around each of the two contained nuclei 
and finally more or less completely segmented off to form two 
formative cells, of which the one placed more internally travels 
to the tip of the spicule ray, while the other remains at the base 

FIGURE 51 SEE 49. 

b.f.c., basal formative cell, tr.syst., triradiate system. 

(Figure 51, b.f.c.}. The apical formative cell sooner or later dis- 
appears, returning, apparently, to the epithelium. The basal 
formative cell remains at the base of the ray (Figure 51), until 
this portion is secreted to its full thickness. It then migrates 
slowly outwards along the ray, and in the fully formed spicule is 
found adherent to the extreme tip." 

The Unity of the Organism 

That such a mode of development is entirely foreign to crystal 
production hardly needs to be remarked. But if further proof 
to the same effect were demanded, one other strong piece of evi- 
dence is the fact that most of the spicules are not composed of 
inorganic substance alone but have a core of organic matter. 

Although the technique for the examination of cytoplasm de- 
veloped during these last years has not, so far as I know, been 
applied to the spicule-producing cells of sponges, we can be 
reasonably sure from the study of other secretory cells what the 
general results will be when such application shall have been 
made. They will bring out numerous details not now known of 
how both the nucleus and the cytoplasm act during spicule pro- 

Surely it is not necessary for me to dwell again on the 
main point of the evidence here presented. The nuclei of 
the spicule-forming cells may take an active part in pro- 
ducing the spicules. Indeed from our general knowledge of 
nuclear activities, illustrations of which were given in an 
earlier chapter, it is probable that such will some day be 
demonstrated to be the case. But the proof of nuclear 
activity in spicule production will not be disproof of the 
already observed cytoplasmic activity in spicule production. 

Other animals that may be mentioned in which spicules 
are produced in much the same way and have the same taxo- 
nomic diversity and constancy are the alcyonaria among 
coelenterates, the holothurians among echinoderms, and 
some of the compound ascidians, particularly of the family 
didemnidae. Relative to the specificity of the structures in 
holothurians, we have this piece of significant information : 
"These calcareous bodies are of great value to the system- 
atist in classifying the smaller groups, such as genera and 
species. Although their general characteristics are fairly 
similar within the several families, the different shapes of 
spicules are not sufficiently constant to be used as diagnos- 
tic characters of such large divisions." 15 In other words, 
so far as these animals are concerned, should it be found, 

Evidence from Somatic Hist ogene sis 


as it almost certainly will be, that the cytoplasm is "hered- 
itary substance" for the production of spicules, the reverse 
of Conklin's generalization that the cytoplasm determines 
the larger taxonomic features of animals while chromatin is 
the seat of the inheritance factors for "filling in details" 
turns out to be true. Species-marking details are just 
what we are able to see the cytoplasm fill in. 

The "Hairs" of Higher Plants 

For a few more instances of species characters in multi- 
cellular organisms brought down to single cells, we turn to 
the plant world. 




The "hairs" or trichomes borne on the leaves, flowers 
and smaller stems of innumerable flowering plants are 
usually composed of only a few cells, so that the characters 
they have are often referable to the individual cells. Can- 
non has lately investigated these structures in several groups 
of plants, and while he was not aiming at the particular 
question now occupying us, some of his results are quite to 
the point for this discussion. An example of particularly 
distinct specificity of the hairs is presented in the following: 
The trichomes of the three species [of poppy] are similar 


56 The Unity of the Organism 

in form and size, but they are unlike in quality of rough- 
ness. In Papaver orientate (52 a, b) and Papaver pilosum 
(53 a, b,) the distal ends of the superficial cells project be- 
yond the general surface of the trichome and turn out at 
a rather acute angle. In Papaver somniferum, however, 
these cells did not extend beyond the general surface, with 
the effect that the trichomes of this species are smooth (fig- 
ure 54 a, b)." 16 

This is especially instructive because the attribute in 
question pertains to the shape and position of cells, and not 
to differentiation within the cells. There, consequently, is 
no room for even a reasonable surmise that the attribute is 
explained by the chromatin instead of the cytoplasm. Even 
though the account gives no information about the position 
and behavior of the nuclei of the cells, it is hardly con- 
ceivable that any one would maintain that the substance 
itself of the cell-tips is not the main factor in the out- 
turning of these tips characteristic of Papaver orientate 
(figure 52 a, b). 

Dealing with quite another type of trichomes, those of 
the walnut, Cannon writes : "A character which easily dis- 
tinguishes the short secreting trichomes of Juglans cali- 
fornica from those of Juglans regia or Juglans nigra 
is the length of the head-cells." 17 Both drawings and tables 
of measurements of the heads showing lengths and diameters 
are given to bring out the positiveness of the distinction. 
Furthermore, details of cell division and cell structure dur- 
ing the development of the trichomes are furnished ; so the 
visible evidence that the cytoplasm is "inheritance material" 
in this case is beyond question. What the invisible evidence 
may be remains for further investigation to discover, but of 
one thing we may be sure : no matter how many facts of 
development now invisible may later become visible they will 
not destroy the validity of the present visible evidence. 

How far it would be possible to go on pointing out specific 

Evidence -from Somatic Hist ogene sis 57 

characters of plants that are referable to individual cells I 
do not know; but judging from the instances that come to 
view even in my limited knowledge of the subject it might 
be carried to an almost indefinite extent. Another instance 
which I recall from the experience of my student days is 
the case of mosses. The serration of the leaves, I remem- 
ber, was one of the features relied upon for generic and 
specific characteristics, and I also remember that the indi- 
vidual teeth often if not always consisted of one or a very 
few cells. 

Cell-wall Structures i/n Higher Plants 

That the cell-wall is a structure of great importance in 
plants is known to everybody ; and the veriest tyro in plant 
histology has learned something of the enormous variety 
and definiteness of character in different tissues and dif- 
ferent kinds of plants presented by this part of the cell. 

A very brief reference to two plant structures, pollen- 
grains and wood tissue, will be, perhaps, a sufficient re- 
minder of what there is for us in this domain. A typically 
formed pollen-grain is a minute spheroidal body containing 
two cells, one known as the antheridial or germinative cell 
and the other as the sterile or vegetative cell. The wall of 
the grain consists of an outer coat, the exine, and an inner, 
the intine. The elaborateness of structure which these coats 
may reach is astonishing if regarded in the crepuscular light 
of the theory that cells are "simple" things. The most dis- 
tinctive thing about the pollen-grain is the pollen tube 
which is produced on one side of the grain and through 
which the antheridial cell reaches the ovule in fertilization. 
The taxonomic variety which is our main interest just here 
pertains largely to the sculpturing of the surface of the 
exine and to the structure of the exine at the point where 
the pollen tube will break through. It is well known to 
botanists that the "spikes, warts, ridges, combs, etc." of 

58 The Unity of the Organism 

the surface of the grains are in general definitive of taxo- 
nomic groups of plants. And concerning the places of 
emergence of the pollen tube we read: "The number of 
these peculiarly organized points of exit is a fixed one in 
each species, and often in whole genera and families." 
As to the way these various structures are produced we have 
this very definite statement : "The sculpturing upon the 
outer surfaces of the spores of mosses and ferns and the 
corresponding pollen grains of the Phanerogams can in 
most cases be attributed to the activity of the protoplasm 
surrounding the developing spores." 

That the main tissues of plants present taxonomic char- 
acters is amply illustrated in the wood-tissues. The facts 
concerning these tissues have been almost forced into 
prominence by the needs of fossil botany, though they have 
also been much studied as a part of ordinary plant mor- 
phology. The section on fossil wood in Zittel's Handbook of 
Palaeontology is a considerable resume of knowledge in this 
field, and contains numerous statements and figures which 
bring out impressively the general truth of the specificity 
of the tissues of trees. After necessary allowance is made 
for the strictly botanical unsatisfactoriness of many of the 
species and genera recognized by palaeobotanists, it is not 
doubted, so far as I know, that on the whole the kinds of 
tissue they describe do in reality represent different kinds 
of trees. A single illustrative quotation will serve to make 
concrete what is here dealt with in general terms : "The 
phloem segments, like those of the xylem, are divided by 
few-seriate pith rays into rather regular two- to four- 
seriate rows of cells made up of thin-walled, small-celled 
elements in the trunk of Zamia floridana, etc., and Stan- 
geria. But ... in the trunks of Cycas, Dion, Encepha- 
lartos, Macrozamia and doubtless most cycads, as likewise 
in the Cycadeoideae, sclerenchymatous elements are more 
or less numerously and regularly interspersed among the 

Evidence from Somatic Histogenesls 59 

row cells, thus giving much more strength to the stem." 20 
And the author goes into considerable detail in discussing 
the presence and the absence of "wood tracheids," "scalari- 
form pittings," "border pits," "spirally thickened walls" 
and "sieve tubes" in the genera Zamia, Cordaites, Stangeria 
and Cycas. 

That this well-nigh endless variety of character of the cell- 
wall in adult plant tissues is due to the activity of the cell 
protoplasm appears never to be questioned by botanists 
so long as they are dealing with the actual structure and 
development of the wall. "The cell-membrane is produced 
by the protoplasm," we read in the section on morphology 
in the LeJirbuch der Botanik by Strasburger, Jost, Schenck 
and Karsten (llth German edition) this section being from 
the pen of Strasburger himself. This simple, unqualified 
statement of fact by Strasburger is the more noteworthy 
because, as we saw in another connection, he has been one 
of the extremists on the chromosome dogma of heredity. 
Reading his statement that the cell-membrane is produced 
by the protoplasm, with the indubitable fact in mind that 
this membrane presents innumerable characters which are 
taxonomically definitive, and hence are hereditary according 
to the best criteria we have of hereditary characters, it 
seems impossible to avoid seeing that it implies an irrec- 
oncilable contradiction of Strasburger's often-repeated 
view that the chromosomes are the sole bearers of heredity. 

To round out the primarily factual part of this discussion 
two questions remain to be considered : first, that of heredity 
in the main classes of tissues of multicellular organisms ; 
and second, that of the results being reached by the latest 
methods of cytoplasmic study on the behavior of different 
portions of the cells in the histogenesis of these tissues. 
What is implied in these two questions can be made clear by 
a special case. Is the minute structure of striated muscle 
tissue, for example, subject to heredity? If so, are the 

60 The Unity of the Organism 

hereditary attributes determined by the chromatin or by 
the cytoplasm of the myogenic cells? Applying our usual 
test for hereditary structures, we ask whether or not this 
tissue presents attributes characteristic of the taxonomic 
groups, species, genera, families and so on. Here again, 
while we have a vast store of knowledge about the structure 
and genesis of muscle tissue in many kinds of animals, only 
incidentally, as a rule, have the studies been made from the 
taxonomic standpoint. 

The Morphology of Striated Muscle Fibers 

For one series of very recent studies that comes near this 
standpoint w r e are indebted to H. E. Jordan. On the taxo- 
nomic aspect of the matter Jordan writes : "A quite gen- 
eral consensus of opinion considers them more or less closely 
related, and ranks them both between Crustaceans and 
Arachnoids. Limulus muscle, however, is in appearance very 
much more like vertebrate than like insect muscle ; while the 
muscle of the marine arthropod Anoplodactylus is of the 
typical insect type." 21 Of the numerous differences between 
the fibers of the two animals compared, reference to two will 
suffice for our purpose. They concern the so-called M 
and Z lines found in the light bands of most striated muscle 
tissue. What Jordan regards as one of the important re- 
sults of his work on Limulus muscle was the evidence secured 
that the Z line represents a membrane, as some observers 
have believed, the specially convincing evidence being the 
fact that the "line" is attached to the sarcolemma periph- 
erally and to the nuclear wall centrally. These rela- 
tions are lacking, he says, in the sea-spider's muscle. The 
M line, he tells us, is especially well developed and hence 
easily demonstrated in the sea-spider muscle in some states 
of contraction, while he failed, as have other students, to 
detect it at all in the Limulus muscle. 22 

Evidence from Somatic Histogenesis 61 

So we come again to the real issue.- Assuming the Z 
membrane to be a cytoplasmic structure, as it has practi- 
cally always been held to be, are w r e going to deny that the 
cytoplasm itself causes the peculiarity of the Z membrane 
in the sea-spider as compared with that in Limulus, that 
denial being necessitated by the dogma that the real "seat' 1 
of the difference is the chromatin of the nucleus operating 
by some invisible "factor" perhaps of the nature of an 

The extent of variety in striated muscle tissue is brought 
impressively to view in such a comparative study as that 
by Marceau. His main object, is not to find differences but 
to discover whether in spite of structural differences they 
have similar traits, as if they might all be derived from a 
single primitive form which has undergone more or less 
profound modification. 

Of the many differences which the investigation sought to re- 
duce to orderliness on the basis indicated, .onlv two will be men- 


tioned. From an elaborate table of measurements of the diameter 
of fibers, we find the following results for the sheep and pig: 23 

Maximum Minimum Average 

Sheep 25 ju 5fj, 15]j, 

Pig 4,5 5 20 

The other point selected concerns "striated scleriform trans- 
verse bands" characteristic of the muscle fibers of the vertebrate 
heart. This time the animals we choose are the horse and cow. 
The thickness of the band is given as exactly the same in these 
two, but the distance between the bands is 140ju for the horse and 
120jU for the cow. 24 

TJw Physiology of Muscle Fibers 

We might go on almost endlessly, pointing out slight but con- 
stant specific differences that involve differences in muscle struc- 
ture, questioning in each case whether the hereditary cause of 
this difference lies in the cytoplasm or chromatin of the muscle 

62 The Unity of the Organism 

cells. But such repetition would be useless for the present dis- 
cussion. Striated muscle tissue is specially favorable for testing 
hypotheses about inheritance material from the functional side 
as well as from the structural side. For example, there are innu- 
merable differences., larger and smaller, in the limb movements 
of animals belonging to different species and genera. I know of 
no observations which precisely connect activities of this sort 
with muscle tissue; but information concerning the electromotor 
force in various animals is available. The following on the au- 
thoritv of Eno-lemann, taken from Winterstein will serve our 

f O 

purpose. The values are those of the "demarcation current" of 
galvanic electricity of heart muscle, this current being generated 
by making a cut surface at the base of the heart and the natural 
surface at the apex act as a galvanic pile: 25 

Animal species Electromotor force in D. 

Anguilla fluviatilis 0.0265 

Rana esculenta 0.0311 

Triton cristatus 0.0124 

Tropidonotus natrix 0.036 

Testudo graeca 0.022 

Columba livia 0.0458 

Cygnus oler 0.0168 

Mus musculus v. albino 0.040 

Mus rattus v. albino 0.0446 

Lepus cuniculus 0.0363 

These investigations appear to have been made from the stand- 
point of general physiology, and therefore not to have been car- 
ried out with the systematic exactness and exhaustiveness de- 
manded for taxonomic discrimination. We may consequently 
presume that more searching examination of the same series 
would considerably modify these results, but we have no reason 
to suppose tint they would eliminate altogether the differences 
due to the animal species. 

After due allowance is made for the purely physiological 
and environic causes which undoubtedly explain a great 
many of these differences, probably no biologist would hesi- 
tate to grant that many of them have an hereditary basis. 
Turning again to the question of the seat of the hereditary 

Evidencr -from Somatic Histogenesis 66 

factors, we are now especially attracted by the functional 
aspect of the subject. To an unsophisticated physiologist 
studying the phenomena involved in this question, it would 
probably never occur that more than one answer is possible. 
Well-informed as such a physiologist may be supposed to be 
on the important part known to be played by the nucleus in 
the life of the cell, he would undoubtedly take it for granted 
that the whole nucleus, its chromatin with the rest, con- 
tributes in some fundamental way to the result. But unless 
well indoctrinated beforehand with the chromosome dogma 
of heredity, he would almost certainly be amazed were 
some one to contend seriously that the cytoplasm is not the 
material basis of the hereditary peculiarities exhibited. He 
would reply, "Why, you are virtually denying that the 
substance of the muscle fiber is the real seat of muscular 
activity, thus implying a contradiction of the 'universally 
accepted principle that the potential chemical energy of the 
muscle substance is the primary source of muscular energy 
in all its manifestations' 26 for surely muscular energy 'in 
all its manifestations' would include those elements of mus- 
cular activity which are hereditarily distinctive of different 
kinds of animals." 

That the cytoplasm is at least the main source of the 
muscle substance furnishing this energy would not be ques- 
tioned, probably, by any histologist, but the definiteness of 
view held at the present time on this subject is worth re- 
calling and is indicated by such statements as the following: 
"The energy of contraction is the transformed surface- 
energy of the ultimate structural elements or colloidal par- 
ticles (submicrons) composing the fibrils." 27 

Presumably there would be much difference of view among 
physiologists as to the validity of the chemico-physical part 
of Lillie's theory of muscular contraction ; but apparently 
there would be very little dissent from that part of his view 
which locates the processes, whatever their exact nature, in 

64 The Unity of the Organism 

the muscle fibers. So it would be merely a matter of suf- 
ficient patience to go over all the tissue systems, epithelial, 
glandular, bony, nervous, and the rest, and point out nu- 
merous certain, and innumerable probable instances of dif- 
ferences for different taxonomic groups of animals, and to 
show that these hereditary differences are expressed pri- 
marily in the cytoplasm of the cells. 

Summary of Positive Information about the Physical Basis 

of Heredity 

We have explored a vast region of fact and theory con- 
cerning propagation and development in organisms, for the 
purpose of ascertaining what is actually known about the 
organs and substances by which hereditary attributes are 
produced. Expressing the matter in terminology familiar 
to current discussion on heredity, we have been trying to 
find what is actually known about the physical basis of 
heredity. If clear-cut, unequivocal information of the 
kind sought is contained in all we have seen, it ought to be 
statable in a few simple sentences. 

What has been accomplished may be epitomized in two 
such sentences: 

First. Overwhelming observational evidence has been se- 
cured that the cytoplasm of cells participates directly in 
the formation of organic parts which have hereditary at- 

Second. A great mass of evidence, partly of observation 
and partly of legitimate inference from the principles of 
organic integration, has been secured, that the chromosomes 
of the germ-cells in plants and animals which propagate by 
means of such cells, participate in the production of or- 
ganic parts having hereditary attributes. 

Any substance which plays such parts in development 
may be named a physical basis of heredity; and these two 

Evidence from Somatic Histogenesis 


groups, or categories of knowledge must, it seems, serve as 
the foundation of all legitimate reasoning about such "basis 
of heredity," or "inheritance material," or "hereditary fac- 
tors," or "bearers of hereditary qualities," or whatever ex- 
pression for the idea be employed. 


1. Payne 190 

2. Meves 816 

3. Meves 817 

4. Meves 820 

5. Meves 850 

6. Meves 845 

7. Cowdry ('14) 416 

8. Cowdry ('16) 436 

9. Lewis and Lewis 393 

10. Loeb ('06) 181 

11. Conklin, ('15) 176 

12. Ritter ('93) 

13. Sedgwick 82 

14. Minchin ('00) 107 

15. Goodrich 

16. Cannon 13 

17. Cannon 23 

18. Goebel 367 

19. Campbell 51 

20. Wieland 197 

21. Jordan ('16) 493 

22. Jordan ('16) fig. 7 496 

23. Marceau 273 

24. Marceau 280 

25. Winterstein Ill 

26. Luciani Ill, 85 

27. Lillie, R. S. ('16) 255 

Chapter XVI 


Antecedents of the Cytoplasmlc and Nuclear Theories of 

Inheritance Material 

FOR the purpose of calling vividly to mind the character 
of the evidence on which, the two propositions rest 
with which the last chapter ended, it will be profitable to 
cast a glance back on the course along which biology has 
come down to us, with a view to finding a sharply out- 
standing spot in the early growth of knowledge which led to 
each of them. On the botanical side such a spot in the 
knowledge of cytoplasm as hereditary substance, is the 
work of Schleiden on the microscopic structure of adult 
and developing plant tissues. The publication of his Ueber 
Phy to genesis, 1838, may be taken as the starting point of 
our knowledge of cellular transformation in the production 
of tissues. It should be remembered that the observers of 
that period had very hazy notions about the distinction 
between nucleus and cell-body, or cytoplasm. On the 
zoological side the publication by Ehrenberg, in 1836, of 
Die Infusionsthierchen als volkommene Organismen may, I 
think, be looked upon as the first milestone in the progress 
of knowledge of cytoplasmic transformation into tissue sub- 

The ever-broadening stream of knowledge of the chromo- 
somes in relation to heredity is usually held to have orig- 
inated in the discovery forty years ago, by O. Hertwig, 


Inheritance Materials of Germ-Cells 67 

that the most essential fact in fertilization is the union of 
the nuclei of the male and female germ-cells. 

That cytoplasm is a physical basis of heredity is proved 
by a great body of direct observational knowledge. That 
the chromatin of chromosomes is a physical basis of heredity 
is proved by much observational knowledge when this knowl- 
edge is supplemented by reasoning involving the principles 
of biological comparison and correlation. These two masses 
of knowledge constitute, as already indicated, the founda- 
tion of all legitimate reasoning about inheritance material. 
Whether chromatin and cytoplasm are the only substances 
which participate in the formation of hereditary structures 
can not now be stated with certainty, though there are both 
observational and general grounds for believing that they 
are not. But into this question we need not enter in this 
discussion. Nor is it necessary for our purpose to inquire 
very particularly whether the conceptions of chromatin 
and cytoplasm really imply just two substances or two great 
classes of substances, though it is best to have in mind the 
undoubted fact that the latter alternative is the true one. 
Beyond a doubt "chromatin" and "cytoplasm" ought al- 
ways to be understood to mean "kinds of chromatin" and 
"kinds of cytoplasm." 

Fwnction of Chromosomes in Heredity Acquired and 


The question which specially concerns us here is that of 
what the relation is between chromatin and cytoplasm in 
virtue of which they play the particular roles they are found 
to play in producing hereditary' structures. Perhaps the 
most important aspect of this general question is that which 
the theory of phyletic evolution naturally brings up: does 
the evidence in hand suggest any answer to the question 
whether chromatin or cytoplasm is the more primitive and 

68 The Unity of the Organism 

fundamental as hereditary substance? Surveying as we 
have the whole field of organic propagation, including the 
process in unicellular organisms as well as in multicellular, 
and sexless as well as sexual methods of reproduction, and 
taking the facts as they actually present themselves, it 
seems as though but one answer to this inquiry is possible: 
The substances included under the generic term cytoplasm 
are the more fundamental and primitive. 

The only possible way of escaping this conclusion is by 
excluding from the conception of heredity the vast majority 
of developmental phenomena presented by unicellular organ- 
isms and by monogenic propagation in multicellular organ- 
isms. As our examination of these provinces revealed, such 
exclusion is exactly what the chromatin dogma of heredity 
has undertaken, implicitly or explicitly, to fix upon biology. 
The utter unwarrantableness of this undertaking was made 
sufficiently obvious, we may assume, by the examination ; 
so we need spend no time on that now. All that is necessary 
is to remind ourselves vividly of the main positive outcome 
of the examination : Heredity is a universal phenomenon of 
the living world. It is coextensive with organic propagation 
and development, while "carrying heredity" by chromo- 
somes is, according to the evidence, very far from a uni- 
versal phenomenon. It is a long way from being coex- 
tensive with organic propagation and development. We 
may, consequently, proceed in our quest of a more rational, 
more consistent, more satisfactory conception of the purely 
operative side of producing hereditary structures. Pur- 
suing the quest, we remind ourselves of having found that 
only the most meager observational evidence is afforded by 
the protophyta and protozoa, and by monogenic metaphyta 
and metazoa, that chromatin is hereditary substance, while 
these organisms afford an overwhelming mass of such evi- 
dence that cytoplasm is hereditary substance. 

But if in the lower, more primitive moiety of the organic 

Inheritance Materials of Germ-Cells 69 

realm, chromatin is a physical basis of heredity to only a 
limited extent and in a partial way but has this office widely 
and positively fixed in the higher moiety, the moiety, that is, 
in which bisexual propagation is fully established, what 
other conclusion can be drawn consistently with the modes 
of reasoning universally sanctioned by evolutionists, than 
that the function of "carrying hereditary qualities" by the 
chromosomes in higher organisms is a secondary or ac- 
quired, or better a delegated or assigned function? Hered- 
ity is far older, phylogenetically, and far broader tax- 
onomically than is the chromatin mechanism by which it 
now in part manifests itself. Under this interpretation the 
acquisition by chromosomes of the function of carrying 
heredity would belong to the same evolutional type as for 
example the acquisition by certain cells of the function of 
muscular contraction, or by certain other cells of con- 
ducting nervous stimuli. The advantage and satisfaction 
of a conception of the role of chromosomes in heredity 
which ranges them naturally and easily with all other or- 
gans and tissues of the plant and animal body will be 
quickly seen by every one to whom the seemingly endless 
chance of discovering new interrelationships and consis- 
tencies in living nature is one of the most rewarding things 
about biological investigation. 

While we are duly impressed with the importance of per- 
ceiving that chromosomes fall into the great class of other 
organs and tissues when considered from the standpoint 
of phyletic differentiation of structure and function, we 
should not fail to notice that within the class they hold on 
a number of counts a very distinct place. Probably the 
most distinctive of these counts, at any rate the one most 
important for this discussion, is the fact that while the vast 
majority of tissues, taking the term in its usual meaning, 
stand at or near the termination of the ontogenic series 
are, in other words, the final stage in the series the chro- 

70 The Unity of the Organism 

mosomes in their function as bearers of heredity stand at 
the beginning. They represent the initial stage in the series. 
Furthermore, their function in this respect is unique as 
contrasted with the function of other tissues, in that while 
other tissues have, typically, each a single function which 
they perform immediately, the chromatin of a given germ- 
cell has a great complex of functions, namely, that of ini- 
tiating the development through which all the attributes of 
the individual during its whole life-career are developed. 
In a word, chromosomes of germ-cells are not determiners or 
carriers of determiners ; they are initiators or carriers of 
initiators. They may, then, be called bearers, or carriers 
of heredity in a very literal sense, namely in the sense that 
they are made use of by one individual, the parent, to carry 
across or transfer from itself to another individual, the 
child, the hereditary attributes of the species in a latent or 
potential state. By virtue of their being thus used, they 
are members of a developmental series, in which series their 
place is at the beginning and not at the end, the nature of 
the series depending on the phylogenic history of the par- 
ticular organism to which the particular chromosomes be- 

The Two-fold Character of the Problem of Hereditary 


At this point it becomes a matter of the greatest im- 
portance for theories about hereditary substance to dis- 
tinguish between the problem of the operation of such sub- 
stance in the developing individual, and that of how such 
substance ever came to be hereditary substance; stated 
otherwise, between the problems of how any substance par- 
ticipates either directly or as an agent in the building up of 
a structure having hereditary attributes, and that of how 
the substance itself became impressed with the attributes, in 

Inheritance Materials of Germ-Cells 71 

a latent state, of the progenitors of the developing in- 

Investigation of the first of these problems is to a large 
extent a matter of observation, as we have seen in the pre- 
ceding pages. The sub-science of histogenesis consists 
largely in tracing out the processes by which completed 
tissues arise by the transformation of less differentiated or 
undifferentiated cells. And when such newly arisen tissues 
and structures are proved to be hereditary by such evi- 
dence as we have called attention to, then the study of 
histogenesis comes to be so far a study of hereditary sub- 

When, however, we turn to the other problem, that of 
how hereditary substance comes to be such, we are in a 
different, a much more difficult case, for so far science has 
succeeded in getting almost no observational hold upon it; 
and despite the vast discussion it has received the darkness 
that envelops it is hardly an iota less black than it was the 
day of its original formulation. But stygian as the darkness 
is, here, especially as to details, we yet are able to see, 
probably, the quarter from which light will come if ever it 
does come. That quarter is the physical-chemistry con- 
ception of the organism as a system of phases the whole of 
which, as a species entity, is essential to its equilibrated 
activities. This nature of the organism, together with some- 
thing akin to its internal secretory and enzymic productiv- 
ity, enables it, we may conjecture, by some means now 
wholly unknown, to reflect its totality of transferable at- 
tributes upon the germinal cells and to transform them into a 
latent state. Ungrudging acknowledgment of the complete- 
ness of our ignorance of how any part of a cell or any other 
portion of an organism becomes endowed with the capacity to 
develop or causally to affect the development of an organism 
similar to that from which it came, should be an important 
item in the preparation to accept any and all indubitable 

The Unity of the Organism 

facts connected with heredity, even though no causal ex- 
planation of them is forthcoming. 

The childlikeness, as Conklin well characterized it, of 
the belief that chromosomes are a simple and complete ex- 
planation of inheritance would not be so bad in itself. If 
it stopped there, as genuine childlikeness would, no posi- 
tive harm could be done. It is the making of this belief 
the starting point of a grand speculation which blinds the 
eyes and closes the mind to a vast number of facts and 
legitimate inferences about heredity, that plays havoc with 
thinking on genetics. It would be a great gain if gen- 
etic theory would recognize wholeheartedly that all or- 
ganic development, as contrasted with mere enlargement, 
consists more fundamentally and obviously in transforma- 
tion of substances than it does in unchanged continuity of 
substances. For under such recognition the futility of 
attempting to explain the transformation of one lot of 
substance by referring it to another lot which does not 
transform, or in other words to explain development by 
something that does not itself develop, would be manifest. 

That the chromosome theory of heredity in reality deep- 
ens rather than illumines the darkness which surrounds the 
problem is seen when one reflects that not only does it 
throw no light on the question of how the chromosomes come 
to be bearers of heredity, but that it creates the new and 
equally difficult question of how the chromosomes (which ac- 
cording to the theory maintain unchanged their individuality 
not only from generation to generation but throughout each 
ontogeny) are yet able to be causally operative in the cell- 
bodies undergoing the transformations which they actually 
do undergo in the developing organism. That the germ- 
plasm-chromosome theory of heredity could have led its 
devotees to sidestep the details of ontogeny, especially those 
of histogenesis, to the extent which our review has shown 
it to have done, would be unbelievable but for what is 

Inheritance Materials of Germ-Cells 73 

actually before us in the recent history of biology. 

Viewing heredity as being definitively a kind of organic 
transformation transformation, that is, in accordance with 
a pre-existing or ancestral pattern more than it is a kind 
of continuity- -it becomes obvious that even were the dem- 
onstration to become complete that the chromosomes are 
the only portions of the germ-cells * essential to fertiliza- 
tion, they still would not be proved bearers of heredity in 
such sense as the germ-plasm theory holds them to be. They 
would not because the problem of the transformations 
which constitute ontogeny would still be untouched. The 
theory would be established only when the demonstration 
should be produced that the chromosomes cause immedi- 
ately all the particular ontogenic transformations known 
to be hereditary. All that would be proved about heredity 
by demonstration that the chromosomes alone participate 
in fertilization would be that the chromosomes alone con- 
stitute the first ontogenic stage of the hereditary parts of 
the particular organism to which the fertilized egg gives 

The Probability That Inheritance Material Becomes Such 

In Each Ontogeny 

But because thus far failure has attended all efforts to 
get knowledge of how hereditary substance is produced, 
are w r e obliged to own that we know nothing at all, even 
inferentially, about its production? And is the search for 
such knowledge to be given up as hopeless? My answer is 
an energetic negative to both these questions. In the first 
place, there is much evidence to support the hypothesis, 
very general to be sure but yet by no means devoid of use- 
fulness, that hereditary substance becomes such in some 

* The utter unwarrantableness of the common assumption that as 
regards the male germ-cell such a demonstration is "practically com- 
plete" will be noticed presently. 

74 The Unity of the Organism 

way through being subject to the metabolic processes com- 
mon to the whole organism. Undoubtedly the germ-plasm 
dogma itself has tended strongly to divert attention from 
this aspect of the problem of germinal material indeed, 
has tended to minimize the importance of the metabolism 
of such material even if it has not tended to deny that the 
material is subject to this process. 

So important is it from the organismal standpoint to 
conceive the material basis of heredity as part and parcel 
of the organism generally, especially as regards the basal 
growth and sustentative processes, that we must examine in 
some fullness the evidence favorable to such a conception. 
In its most brazenly evidence-ignoring form, the germ-plasm 
dogma asserts that the female parent does not really pro- 
duce the eggs or the male parent the sperm, as they seem 
to, but that these are produced by previous germs ad in- 
finitum. There are, to be sure, quite a number of observ- 
able facts, as those of the early formation of germ-cells 
in several animals, that can be forced into a seeming sup- 
port of such a conception. But the familiar and all but uni- 
versal fact that multicellular organisms, plants and animals 
alike, are sexually immature for a shorter or longer part of 
their lives, the very essence of the immaturity being the un- 
developed state of the reproductive system, would be a 
sufficient refutation of the view for any mind not made 
impervious to facts by long and faithful sophistication. 

Germ-Cells Subject to Metabolism Like All Other Cells 

The biological commonplace that all germ-cells, like all 
other cells, undergo a process of growing and maturing 
before they can perform their distinctive office, and that 
this process depends upon the retention of the germs by the 
parent organism, ought, as already indicated, to be a suf- 
ficient antidote againjst the germinal continuity fallacy, 

Inheritance Materials of Germ-Cells 75 

even though nothing were known as to exactly what goes 
on in the germ while it is growing and ripening. But we 
are by no means without positive knowledge under this 
head. In fact the last few cell divisions immediately pre- 
ceding the ripening of both ova and spermatozoa, and the 
ripening processes themselves, have received searching ex- 
amination during the last few decades, with the result that 
hardly any cytological phenomena are better known than 
are the profound morphological changes which accompany 
these processes. That these changes are particularly mani- 
fest in the chromosomes, the assumed seat of the determiners 
of heredity, is one of the very things that has aroused so 
much interest in the processes. Nor are we wholly unin- 
formed about the chemical changes taking place in the 
growing germ-cells. Unfortunately knowledge in this field 
has hardly passed the stage of early infancy, but at least 
enough is known to warrant the assertion that the young 
germ-cells are subject, as are all the other cells, to the 
general metabolism of the organism. 

Chemical Changes in Germ-Cells During Parent's Ontogeny 

About the most striking information we have in this 
field is what has come from such investigations as those 
on the chemical changes which occur in the sex glands and 
other body parts during reproduction in some fishes.* Mie- 
scher's work was ground-breaking in this domain for it 
was the first to show that the "sexual organs in the salmon 
develop at the expense of the muscular system, and that the 
salmine deposited in the testis during the breeding season 
must be derived from the proteins of the muscle, since the 

* Notable among these studies are: Histochemische und physiologische 
Arbeit en, gescvnimelt und liermisgegeben von seinen Freunden, by Mie- 
cher; and Changes in the Chemical Composition of the Herring during 
the Reproduction Period, by Milroy. Biochemical Journ., v. in, 1908, p. 

76 The Unity of the Organism 

fish does not take any food during the period." The work 
of Riddle and his collaborators is producing evidence to the 
same effect. 

Such researches do not, to be sure, prove that the chem- 
ico-physiological changes extend to the chromosomes of the 
germ-cells, much less to the imaginary determiners of hered- 
ity in the chromosomes. But viewing the results in the 
light of the well-grounded general belief that the moist 
fundamental test of living substance is metabolic change, 
it is seen that any hypothesis which assumes the existence 
in the germ-cells of something virtually not subject to the 
general metabolism of the cells, assumes at the same time the 
burden of furnishing objective evidence that such a something 
does exist. 

As a matter of fact the prime offense of the germ-plasm- 
determiner hypothesis is that its very essence places it 
beyond the reach of scientific observation. Such truth as it 
may contain cannot be made really effective because it can 
not be proved, and such error as it may contain can not 
be robbed of its power for evil because it cannot be dis- 
proved. In a word, the hypothesis is one that belongs to 
the realm of dialectics primarily, and has no just claim to 
a place in inductive science. 

The Possibility of Changing Sex By Influences on the Germ 

But perhaps the most conclusive evidence of the funda- 
mental dependence of true germinal material upon the or- 
ganism, should somewhat fuller verification of the obser- 
vations be obtained, are results like those reached by Whit- 
man, King, Whitney, Riddle, and by R. Hertwig and his 
students, according to which sex may be reversed in several 
animal species by various conditions extraneous to the 
germ itself, acting on the germ-cells from which the animal 
is to develop. The instance of this usually regarded as best 
established is afforded by certain species of frogs and toads. 

Inheritance Materials of Germ-Cells 77 

The widely known result reached by Hertwig and verified 
and extended by Kuschakewitsch expressed in a single sen- 
tence, is that the number of males and of females produced 
by the eggs of a given female depends upon whether the 
eggs are fertilized when newly ripe or when over-ripe, a great 
predominance of males coming from the latter class. In 
a tabulation of the results of four sets of experiments by 
Hertwig 2 almost every case shows the number of males 
increased when the time elapsing between deposition of the 
eggs and fertilization was increased, the highest percentage 
of males in any one lot being 759, fertilization in this in- 
stance having been twenty-two hours after the fertilization 
of the last preceding lot. 

In a species of toad, Bufo lentiginosus, results have been 
obtained just the reverse of those on the frog, that is, the 
proportional number of females has been experimentally 
increased. This was accomplished by fertilizing the eggs 
in water made slightly alkaline. Since frog eggs are known 
to absorb water when they remain long in it, as in the case 
of those which gave rise to a preponderance of males in 
the Hertwig method of experimenting; and since alkaline 
solutions extract water from eggs, and likewise cause them 
to produce a preponderance of females, Miss King drew 
the obvious conclusion that the quantity of water contained 
in eggs of these animals may be a factor in determining the 
sex of the animals developed from the eggs. Although 
Miss King recognizes that her experiments do not furnish 
final proof of the conclusion she draws, she believes, rightly 
it would seem, that they weigh heavily in that direction. 
"As they stand," she writes, "the results strongly suggest 
that sex in Bufo is determined at or near the time of fer- 
tilization, and that external factors acting during this pe- 
riod may influence the sex-determining mechanism in such 
a way as to cause it to produce one sex or the other. The 
results also seem to indicate that in Bufo sex is determined in 

78 The Unity of the Organism 

the egg, and that it may depend in some way on the rela- 
tive amount of water in the egg at the time of fertiliza- 
tion." 3 

Riddle, perhaps the most outspoken opponent of sex pre- 
destination now writing, strongly espouses the hypothesis 
that the sex to which a particular egg will give rise is de- 
pendent partly on the quantity of water which that egg 
contains. But whether water is a factor in determining sex 
or not, the evidence presented by Riddle, coming partly from 
researches by C. O. Whitman and partly from his own, 
constitutes, when taken with the evidence to the same effect 
presented by other investigators, almost if not quite com- 
plete proof that sex is not the hard-and-fast thing which 
most present-day genetic speculation would make it. 

Furthermore the evidence produced by these two inves- 
tigators seems to connect the decision as to which sex a 
particular egg shall give rise, with some condition of the 
parents. It is well known to all zoologists, in the United 
States at least, that at the time of his death Professor Whit- 
man had accumulated a vast store of data on the habits, 
particularly the breeding habits, of pigeons. To Doctor 
Riddle, who had worked with Whitman considerably, fell 
the task of carrying on to some extent Whitman's experi- 
ments and of preparing for publication the results which 
Whitman left in the rough. The following quotation from 
Riddle's paper referred to above, summarizes Whitman's 
results that are especially important for us now: "Whit- 
man found that if certain very distantly related pigeons 
[i.e., two individuals from different families] are mated 
that only male offspring resulted. If the matings were 
made of individuals not quite so distantly related different 
genera usually and if to this situation be added the ele- 
ment of overwork at reproduction [i.e., the birds not being 
permitted to nest their own eggs, but forced to keep laying 
eggs in rapid succession] then the first several pairs of 

Inheritance Materials of Germ-Cells 79 

eggs produced in the spring will produce all or nearly all 
males. The last several pairs of eggs laid in autumn will 
produce all, or nearly all, females. At the transition period 
in the summer he found that some pairs, or clutches, of eggs 
produced both a male and a female. In these cases it was 
usually the first egg that produced the male ; and the sec- 
ond egg laid forty hours after the first that gave rise 
to a female." 4 

Into Riddle's interesting discussion of Whitman's results 
and his own chemical studies on the eggs of pigeons and 
hens we need not go. Suffice it to say that it seems to me 
Riddle is justified by the evidence now in our possession, 
in his contention that "sex rests upon a quantitative and 
reversible basis" and that in this sense it has been controlled 
by conditions extraneous to the germ-cells themselves. This 
does not imply, as I understand, that such control would 
necessarily be practicable or even possible in all organisms, 
nor does it preclude the possibility that in some species 
there may be dimorphic or partially dimorphic spermatozoa 
or ova as regards sex production. Neither does it preclude 
the possibility that in some cases where a preponderance 
of one sex has been observed, this is due to selective mor- 
tality or some process other than the actual shifting of the 
sex tendency in the particular eggs. 

These several concordant bodies of testimony must, it 
would appear, open the eyes of biologists sooner or later to 
the ludicrousness of a theory that would make the parent 
organism hardly more than a combined culture medium and 
incubating oven for its germ-cells. 

The Determiner Conception Contrary to Ordinary Chem- 
ical Principles 

If, on the basis of such facts as we have, we try to come 
still closer to the questions of how the assimilative and 
morphogenic processes of the organism occur, whether in 

80 The Unity of the Organism 

the production of hereditary substance or in the transform- 
ation of such substance into actual hereditary structures 
and activities, we find ourselves hedged about on every side 
by partial knowledge, by dubious knowledge, and by com- 
plete ignorance on many fundamental points. 

However, chemical considerations seem to point the way 
to future discovery. In the first place, it seems necessary 
to recognize that the whole germ-plasm conception as orig- 
inally promulgated, with its interminable system of "bear- 
ers" and consummators, was contrary to what is well known 
about chemical processes generally. Thus the continuity 
presented by a complex chemical operation does not consist 
in an unchanged series of individual entities of some sort, 
such as determinants and determiners are, or originally 
were conceived to be, but rather in a regular succession of 
transformations. For example, when chromic hydroxide, 
which is grayish-green, is dissolved in acid, a green solu- 
tion results, which turns to greenish violet or pure violet 
if allowed to stand a long time. Exactly what the chemical 
changes are that correspond to these color changes I do 
not know, and probably the information which chemists have 
on the subject is not exhaustive. At any rate modern 
chemistry conceives the phenomenon to consist in a succes- 
sion of reactions and transformations, the various colors 
and shades being attributes of the compounds that exist 
in the various stages along the way, and not as due to in- 
dividual bodies carried by the preceding stages for the 
express and exclusive purpose of producing the particular 
colors that are observed, as would be implied in such a 
metaphysical scheme as was the germ-plasm theory elab- 
orated by Weismann. 

Enzymic chemical action presents perhaps a still better 
starting point for imagining what the fundamental hered- 
itary processes may be than does ordinary chemical activity 
like that just instanced. The essence of this kind of activ- 

Inheritance Materials of Germ-Cells 81 

ity is, as everybody knows, that in some way enzymes cause 
or at least facilitate transformation in other substances. 
Thus the attribute of solubility of the sugar into which 
starch is transformed, through the action of the salivary 
enzyme ptyalin, is not held to be due to a determiner for 
solubility carried by the ptyalin and passed 011 into the 
sugar, but rather it is recognized that solubility is one of 
the attributes possessed by the kind of sugar into which 
starch is converted by the ptyalin. The solubility is thought 
of rather as an attribute of the sugar and not as something 
once latent in the ptyalin which produced the sugar. A 
few details of the action of the enzyme in this case illustrate 
the point still better. Maltose, which is the chief if not 
the only sugar resulting from the action of ptyalin, is not 
reached by a single bound, as one might say, but. through a 
series of bodies known as dextrins, at least three of which 
have been recognized. These are amylo- erythro- and 
achroo-dextrin, named from the color they display when 
treated with iodine, the first mentioned turning blue, the 
second red, and the third remaining colorless. What mod- 
ern chemist would think of explaining the blue of the amylo- 
dextrin by a "determiner" for that color in the ptyalin 
or even in the starch, the red of the erythrodextrin by an- 
other determiner for red, and so on? That is the sort of 
explaining chemists of a century ago did, but they have 
long since learned not merely the futility but the scientific 
evil of such explanation. 

If it were germane to our present task we might go on 
and show that the gene conception in modern genetics is 
really a revival in biology to-day of the gene conception 
which passed muster in chemistry a hundred years ago, 
when oxygen and hydrogen were named. Such an exposi- 
tion would be appropriate to a history of scientific theory 
or to a treatise on the theory of natural knowledge, but 
hardly to the present work. 

82 The Unity of the Organism 

Endorsement of E. B. Wilson's Proposal to Drop "De- 
terminer" From the Vocabulary of Genetics 

In his Croonian Lecture having 1 the title The Bearing of 
Cytological Research on Heredity., E. B. Wilson said, "In 
the meantime it would be well to drop the term 'determiner' 
or 'determining factor' from the vocabulary of both cytol- 
ogy and genetics." If the facts and arguments set forth 
in the preceding pages are valid, they constitute a demon- 
stration that not only would it "be well to drop the term 
'determiner,' but that it must be dropped, at least in its 
present application, before thought and investigation on the 
mechanism of heredity can be free and in very deed truth- 
seeking. "What we really mean to say," Wilson continues, 
"is 'differential' or 'differential factor,' for it has become 
entirely clear that every so-called unit character is pro- 
duced by the cooperation of a multitude of determining 
causes." So far as these statements go they are in strict 
accord with the organismal standpoint maintained in this 
Volume, and we may also say, with the physical-chemistry 

Where attributes of adult organisms have been so defin- 
itely correlated with particular chromosomes and possibly 
parts of chromosomes of the germ-cells as seems to be the 
case in the fruit flies, such chromosomes are unquestionably 
differential, and since they stand at the very beginning of a 
long and complex transforming and developing series, they 
may very properly be called differential factors even though 
they do not themselves participate suhstantively in the 
transformation. The general similarity of their mode of 
action to that of enzymes is certainly considerable : a minute 
quantity of the substance is capable of inducing or facili- 
tating the transformation of a large amount of other sub- 
stance in a perfectly definite manner, and the inducing agent 
is not itself consumed. 

Inheritance Materials of Germ-Cells 83 

Advantages of Conceiting Germ-Cell Chromosomes as Initia- 
tors in Hereditary Development 

This chemical way of viewing chromatin and chromosomes 
sanctions the idea that these are to be regarded as initiators 
of developmental processes which lead to hereditary attri- 
butes, rather than determiners of those attributes. If one 
wants to know in what way this conception would have 
an advantage over the determiner conception as a working 
hypothesis, my reply is that the advantage is two-fold. 
First, it would surely correct the tendency of genetic re- 
search under the guidance of the determiner hypothesis, to 
restrict its attention to attributes of adults at one end of 
the ontogenic series and to the chromosomes of the germ- 
cells at the other end, and to ignore or touch only in the 
lightest way all the intervening parts of the series. This 
correction would result because the new standpoint would 
bring the whole series of continuities and transformations 
alike into proper perspective, revealing thus that the mem- 
bers of the series intervening between germ and adult must 
be investigated in exactly the same way and with the same 
objects in view as the end members, if complete understand- 
ing of the hereditary process be the goal of research. It 
could not then happen that the egg-cell would be repre- 
sented, as it now so commonly is, as a relatively large sac 
containing nothing significant for heredity except the rela- 
tively small chromosomes. Nor could nearly the w r hole 
mass of ontogenic phenomena, especially those of histo- 
genesis, be treated so lightly in speculation and so largely 
neglected in investigation as they have been under the 
domination of the determiner theory. 

The second advantage in the initiator conception is that 
since it would recognize the "differential factor" of the 
chromosomes to be in reality due to the fact that the whole 
ontogenic series to which the chromosomes belong is differ- 

84 The Unity of the Organism 

ential, that is, that it pertains to a particular species or 
kind of organism, it would put an end to the notion by 
which recent genetic science has been so largely dominated, 
that the problem of how the series came to be thus specific 
or differential may be solved by speculation, and it would 
incite geneticists to efforts to solve the problem by obser- 
vation aided by experiment. It is impossible to refute the 
charge that genetics is to-day more interested in an elab- 
orate system of conceptions of speculation, in other words 
than it is in observed or possibly observable phenomena. 
We cannot keep too constantly before our minds the fact 
of our almost complete ignorance of how any substance 
becomes hereditary substance whether through the "inheri- 
tance of acquired characters" or in any other way. Hence 
mere speculation on the subject after the manner of the pan- 
gens idea is much worse than nothing if permitted to run into 
a bewildering and enslaving system like that of the germ- 
plasm theory as it came from Weismann's mind. Neverthe- 
less it is quite germane to the present discussion to point 
out that whatever might be the nature of the chemical ac- 
tion, whether enzymic or some other, through which the 
series of ontogenic transformations should be accomplished, 
the character and subtlety of these processes seem to make 
them, more than any others we know, competent with some 
modification to serve as the go-between for impressing the 
germinal material with the latent attributes of the species. 

Inconclusweness of the C Biological Evidence Usually Ap- 
pealed to in Support of the Chromosome Theory 

And this leads to the concluding statements of this dis- 
cussion. The three categories of cytological fact which 
have been weightiest in the formation and maintenance of 
the chromosome theory of heredity are the individuality and 
continuity, chiefly numerical, of the chromosomes from par- 

Inheritance Materials of Germ-Cells 85 

ent to offspring; the apparent equality (it should never be 
forgotten that the dogma of equality does not rest on rigor- 
ous quantitative investigation) of the chromosomes in the 
male and female germ-cells ; and the assumption that in 
the male germ-cells the chromosomes alone are concerned 
in fertilization. Even if these groups of assumed fact were 
established with absolute certainty they would fah 1 far short 
of being direct and final proof that chromatin is the only 
hereditary substance. That this is true must be apparent 
to all well-informed, carefully thinking biologists. The 
most important grounds for this inconclusiveness are in- 
volved in the facts and arguments set forth in the preced- 
ing pages, but they may be summarized here, and in addi- 
tion two other grounds may be pointed out. 

First and foremost, in my opinion, is the general truth 
that chromosomes or even chromatic substance can not pos- 
sibly be recognized as the sole bearers of hereditary sub- 
stance, because the evidence is enormous in quantity and 
direct and indisputable in quality that other substances par- 
ticipate actively in the production of hereditary attributes. 
There is no way of escaping this conclusion except by nar- 
rowing the definition of heredity for the very purpose of 
bringing it within the scope of the chromosome theory of 
hereditary substance. The scope and fundamentally of 
this aspect of the problem is sufficiently dwelt upon, we 
may hope, in what has gone before. 

The two additional grounds for skepticism as to the con- 
clusiveness of the cytological evidence will now be pointed 
out. First, as to the evidence from the individuality and 
continuity of the chromosomes. All that any careful 
thinker claims or can claim for this evidence is that the 
individuality and continuity of chromosomes as observed 
are what might be expected if they were the germinal de- 
pository of the organism's hereditary attributes. And the 
question is constantly and naturally asked, what other 

86 The Unity of tlie Organism 

meaning can the whole remarkable series of phenomena of 
maturation and fertilization have than that attributed to 
them by the chromosome theory? My rejoinder to this 
argument may begin with a reply to the question. As long 
as knowledge of the chemistry and physiology of cells 
germ-cells with the rest- -is as fragmentary and inconclusive 
as it now is, certainty as to the meaning of the phenomena 
mentioned is out of the question. However, it would seem 
quite probable that they are concerned primarily with the 
nutritive and assimilative processes of the cell and only 
derivatively with heredity. Furthermore, the question asked 
may weU be paired off with another of similar character, 
namely, what is the meaning of the almost if not quite com- 
plete breaking up and disappearance of the chromosomes in 
the so-called resting stages of the immature germ-cells, this 
being accompanied by the dissolution of the nuclear mem- 
brane so as to allow the freest possible commingling of the 
whole nuclear contents with the cytoplasm of the cell? Have 
we not at least as much factual right to suppose that during 
this mixing of substances the chromatin, and so later the 
chromosomes, are influenced by the cytoplasm, as that the 
reverse influence takes place? Is it not entirely possible 
that this process is one of the very means or occasions of 
impressing the chromosomes with the attributes of the or- 
ganism which, as we have seen, apparently must take place 
whether acquired characters are ever inherited or not? 

And now as to the argument from the assumed exclusive 
participation of the chromosomes of the male germ-cell in 
fertilization. First of all, it can not be admitted for a 
moment that the chromosomes are proved to be as ex- 
clusively the fertilizing agents as they are generally as- 
sumed to be. Even in such extreme cases of seeming ex- 
clusiveness of chromosomal participation as that claimed 
by Strasburger for the pollen cells of some plants, neither 
Strasburger nor any one else has claimed, so far as I know, 

Inheritance Materials of Germ-Cells 87 

that all other male substances than chromatin are excluded 
as rigidly as would be required by experiments in a chemical 
laboratory designed to ascertain the action of a particular 
chemical element or substance in its purity. It is certain, 
for example, that in almost if not quite all male germinal 
elements in animals, a thin outer layer from the cytoplasmic 
part of the spermatic! is present on the head of the sperma- 
tozoon. Furthermore, it is well known that at least the 
"intermediate piece" of the sperm tail, which is not usually 
regarded as chromosomal in origin, remains in the egg at 
fertilization. Nor is there any good ground for supposing 
that the non-chromatinic portions of the nucleus are ab- 
solutely excluded. The almost certain presence in the egg 
at fertilization of at least these male substances other than 
chromatin can by no means be regarded as insignificant for 
heredity, especially if the initiator conception of germinal 
material is held. It seems to follow of necessity that if the 
fertilizing substances, whatever their source, be conceived 
to act in an organic system of the physical chemistry sort 
after the manner of enzymes, no such quantitative relation 
subsists between these fertilizing substances and the prod- 
ucts of organic growth as the chromosome theory implies ; 
nor can their action be so narrowly localized in the eg-g-. 

/ oo 

Their action would be conceived to involve the entire ovum 
ab initio, and not the chromosomes alone. 

Summing Up of the Findings Against the Chromosome 


The general result of our critique is that the whole at- 
tempt to interpret the physical basis of heredity in accord- 
ance with elementalist conceptions has failed and must con- 
tinue to fail, so far as its main aim is concerned. We are 
led to see that the germ-plasm dogma, no matter how often 
or how completely it changes its nomenclatural habiliments, 

88 The Unity of the Organism 

as in the shifting from determinants to determiners, or from 
determiners to gem, or from gens to factors, involves a 
rejection of the conception that the germinal elements of 
organisms after being discharged are literally detached 
parts of those organisms. This conception was well on the 
road to incorporation into the great body of established 
biological truth when it was headed off by Weismann's 
diametrically opposed hypothesis of germinal isolation. 

I would insist that the defense of the organismal concep- 
tion in this volume is really a carrying out of such a con- 
ception of the organism and its germinal products as is 
implied by the old view that the germ is a part of the parent 
organism. It would hardly be possible to express more sat- 
isfactorily in a single sentence the most inclusive theo- 
rem, as it might be called, the demonstration of which is the 
aim of the part of this volume devoted to the means by which 
organisms propagate their kind, than the following from 
E. B. Wilson : "To the modern student the germ is, in 
Huxley's words, simply a detached living portion of the 
substance of a preexisting living body carrying with it a 
definite structural organization characteristic of the 
species." 6 

Coupling this statement by Wilson with another from 
one of his latest writings, 5 to the effect that we ought to 
drop the term determiner because in reality what it means 
is differential, I call attention to the fact that the "dif- 
ferential factor" of the later statement and the "definite 
structural organization characteristic of the species" of 
the earlier statement are in essence one and the same. The 
only difference is that in the earlier statement it is the 
whole germ-cell that is recognized to be a detached part of 
the organism, while the later statement can be brought 
down to the chromosomes because of the greater refinement 
of knowledge attained since the earlier one was made. The 
point I wish to make stand forth with the greatest possible 

Inheritance Materials of Germ-Cells 89 

boldness is that the germ-cell chromosomes may properly 
enough be said to be differential, if only one never loses 
sight of the fact that they are differential in no other sense 
than are any other particles or substances of the germ- 
cells or any other cells which participate in the production 
of species-attributes. 

Brief Reference to the Untoward Implications of the Germ- 
plasm Conception of Heredity 

The somewhat laborious task of exhibiting the difference 
between conceiving the physical basis of heredity from the 
elementalist and from the organismalist standpoints may 
well be brought to a close by calling attention to the impli- 
cation of the two conceptions as applied to heredity in man 
himself. Looked at from this direction the germ-plasm 
dogma is seen to be chargable with the grave offence of 
having added its weight to a conception of human life, the 
overcoming of which has been consciously or unconsciously 
man's aim throughout the whole vast drama of his hard, 
slow progress from lower to higher levels of civilization- 
the conception that his life is the result of forces against 
which his aspirations and efforts are impotent. As ap- 
plied to man this form of fatalism is no less sure and no less 
dire in its tendencies than have been any of the innumerable 
theistic forms of fatalism that have prevailed through the 
centuries. It is almost certain that the ardor with which 
Eugenics has been espoused by several biologists is due 
in considerable measure to the fact that they have felt 
more or less definitely this sinister implication of the theory, 
and have turned to Eugenics as the only weapon against 
its evil forebodings. The germ-plasmic eugenist virtually 
says, "Yes, indeed is man a reasoning, willing, aspiring 
animal, but all his activities in these ways are futile so far 
as the race as a whole is concerned, except as they are 

90 The Unity of the Organism 

brought to bear, extrinsically and operatively rather than 
organically, on the Germ-Plasm." This form of the Eu- 
genic idea corresponds in spirit to the propitiative offerings 
of primitive religion. It aims to mollify by human agency 
powers that act upon men's lives, but which are in them- 
selves largely extraneous, largely evil, and wholly irre- 

What eugenists of this school have failed to see, evidently, 
is that even were unit-factors as differentiate from one 
another in heredity as the extremest Mendelist conceives 
them to be, and that even were the germ-plasm improved up 
to the level of his highest hopes, his results in terms of ac- 
tual human lives and social conditions would be distressingly 
meager. They would be so, because whether unit-factors 
exist independently in heredity or not, they certainly do 
not exist thus independently in development and function. 
In these ways they interact upon one another in the most 
vital manner, as physiology, especially of the internal se- 
cretions and the nervous system, and as physiological and 
social psychology are rapidly and conclusively demon- 

We thus end our examination of the means by which or- 
ganisms produce others of their kind with the conclusion 
that the material through which reproduction is accom- 
plished is in the most vital way part and parcel of the 
organism, that is, that the germ-cells are somehow stamped 
through and through, potentially, with the characteristics 
of the kind, or race, or species to which the producing or- 
ganism belongs. And with this we are ready to pass to the 
examination of those integrative phenomena of the organ- 
ism generally, one manifestation of which is this very nature 
of the germ-cells. 


1. Marshall 292 4. Riddle 10 

2. Hertwig, R. ('12) 75 5. Wilson, E. B. ('14) 351 

3. King 232 6. Wilson, E. B. ('00) 7 




Chapter XVII 

The Field to be Covered by the Constructive Discussion 

A CCEPTING the inevitable destructive result of our cri- 
^~JL tique of the elementalist standpoint, that the attempt 
to interpret living beings in the terms of their constituent 
parts alone always leads to partial failure and disappoint- 
ment, or to the worse result of illusionment as to the trust- 
worthiness of the explanations proposed; and accepting the 
constructive result that everything in the critical study tends 
to show that no part of any organism can be rightly inter- 
preted except as part of an individual organism, this indi- 
vidual being in turn interpreted as a member of a taxonomic 
group, it is revealed that we are only on the threshold of the 
positive, the constructive side of our general enterprise. Even 
though the conclusion be unescapable that the living organ- 
ism somehow acts causally on its parts, the problem still 
remains as to the modus operandi of that acting. The "some- 
how" which came to us as an incident of our critical study 
has yet to be inquired into. 

Stated more specifically the task now before us is that of 
examining closely and systematically the interdependences 
among the parts of the individual organism. Although these 
interdependences are among the most obvious and general 
of all organic phenomena such an examination of them biol- 
ogy has not yet made systematically. Indeed and here is 
one of the most vital things for us to see a cardinal charge 
against the elementalist standpoint is that in its very nature 


94 The Unity of the Organism 

it not only does not encourage, it actually stands against 
such examination. Its opposition to comprehensiveness and 
systematization is profound and essential. 

Our examination will begin with a single brief, two-parted 
definition : The structural and functional interdependence 
found to exist among the parts of an organism we call bio- 
integratedness; and the process of moving on from grade 
to grade of interdependence among the differentiating parts 
which constitutes ontogenesis in the individual we call bio- 

Four Types of Bio-integration to Be Treated 

In the present state of knowledge and for the discussion 
now before us four types or kinds of bio-integratedness and 
bio-integration may be recognized as pertaining to the in- 
dividual organism : 

1. Growth integration, most obvious in graded meristic 
series, but also expressed in the "axial gradients" of Child. 

2. Chemico-functional integration, known so far chiefly 
in connection with internal secretions. 

3. Neural integration, comprising the interdependences 
among the parts of the nervous system, and the involvement 
with this of the muscular, glandular and other organs. 

4. Psychic integration, very closely connected with neural 
integration, but approached from the side of the totality of 
activities of living beings rather than from the side of nerve- 
organ activity, and so taking cognizance of a vast number 
of phenomena not yet definitely correlatable with neural 

The full presentation of facts and arguments under these 
four heads would reach far beyond the limits set for the 
present work. We are, consequently, obliged to restrict 
ourselves to a small portion of the best established and most 
compelling evidence under each head. 

Growth Integration 95 

Graded Repetitive Series as Integrative Phenomena 

This, perhaps the simplest form of integrational phenom- 
ena known to biology, is seen "almost everywhere, but shows 
itself most typically and strikingly in plants and in many 
lower animals. Reference is made to the gradation in the 
repetitive or meristic parts appearing in so many organisms. 
The most obvious criterion of such gradation is the rela- 
tive size of the parts, but, as we shall see later, there is con- 
siderable reason for supposing the gradation is not re- 
stricted to size. The few examples to which space can be 
given are selected to represent as wide a range as possible 
of the phenomena under consideration. 

Illustrations from Animals 

The lancelets, fish-like animals of the genus Amphioxws, 
may be noticed first (Figure 55). It will be observed that 


notochord. cir., cirri, or.hd., oral hood, myom., myomeres., 
dorsal fin rays, cd.f., caudal fin. gon., gonads 

the creature tapers off toward both ends and that the series 
of metameres, myom, usually called myomeres because they 
compose the main body-musculature, diminish not only in 
dorso-ventral measurement from near the middle each way, 
but are also thickest in the mid-region and become thinner 
as they progress toward each end. 
Something of this size scheme of body-parts is very com- 

96 The Unity of the Organism 

mon in the animal kingdom. Figure 56, a photograph of 
the skeleton of a python (see frontispiece), shows in a gen- 
eral way the size-relations of metameric skeletal parts in a 
higher vertebrate. Something of the extent to which the 
proportionality of parts of the individual metameres is car- 
ried out in this skeleton is shown by tabulating a series of 
measurements of the parts : 

Position of 


L.V. T.V. T.ex.zyg. H.D.S. W.D.S. 



6 mm. 

5 mm. 

1 1 mm. 

15 mm. 

4 mm. 

25 mm. 
















































"Position of vertebra" refers to the serial number, beginning 
with the head-end, of the vertebra measured. Legend: L.V., 
Length of vertebra, measured from the posterior edge of one 
dorsal spine to the anterior edge of the one next behind it. T.V., 
thickness of vertebral centrum in its thinnest part, i.e., near the 
middle. T. ex. zyg., thickness of vertebra at extreme of posterior 
zygopophyses (articulating processes). H.D.S., height of dorsal 
spine. W.D.S., width of dorsal spine. L.R., length of rib. 

The starfishes are another class of animals which exhibit 
beautifully this size gradation of repeated parts, both their 
"tube-feet" and the calcareous skeletal supports being 
graded proportionately to the tapering arms of the animal. 
The following table presents a series of measurements of the 
two organ systems just mentioned, from a single arm of 
Astrospecten calif ornicus. The dimensions are in millimet- 
ers, and the series proceed from the proximal to the distal 
end of the arm. 

Growth Integration 97 


Series Tube-feet Ambulacral plates Adambulacral plates 

number Length Length Width Thickness Length Width Thickness 

























































































Not only do these graded meristic series appear in the 
individual makeup of a great range of animal species, but 
they occur in the colonies of many species in which aggrega- 
tions are produced by budding. Sometimes, as in many al- 
cvonaria, the size gradations are very obvious, while in other 
groups the distinctions are so small as to be discoverable 
only by close quantitative study. An example of this latter is 
furnished by the plumularian hydroids. A typical colony of 
the genus here studied, Torrey writes, "closely resembles a 
feather, of which the shaft is represented by the stem and the 
veins by the two ranks of alternating branchlets, or hydro- 
cladia, corresponding to barbs. Each hydrocladium is 
divided by more or less definite nodes into internodes and 
bears on one aspect- -the same in all hydrocladia a compact 
series of hydranths, one to each internode." 

Without entering into the tabular and graphic details 
contained in this study, the author's summarized statement 
concerning one of the tables will suffice: "It will be seen from 
the table that, as the tip of the colony is approached, not 


The Unity of the Organism 

only do the hydrocladia possess fewer and fewer hydroth- 
ecae, but the dimensions of the latter through the mesial 
nematophore reaches its minimum more and more rapidly. 
Since the hydrothecae, once formed, do not enlarge with age, 
it is clear that for such colonies as this, there is a limit of 
growth and a specific form." 

This correlation and proportionality among repetitive 
parts is frequently observed within the bounds of particular 


a, axis, a', axis of branch, b', b", primary and secondary branches. 

organs, as for example, in the branching tentacles occurring 
in various groups. A detailed study of one of these cases 
was made by me some years ago on the tentacles of an as- 
cidian. Figure (pi. 1, figure 13) of this study, sup- 
plemented by the following statement, illustrates the 
point. "Although this figure is diagrammatic in a way, it is 
accurate as to numbers of branches. The positions, too, of 
all the branches and length of the primaries were deter- 
mined by micrometer measurements, and the secondaries were 
drawn as accurately as possible." 2 For the rest, the figure 
(Figure 57) tells its own story. 

Growth Integration 


Illustrations from Plants 

But it is in the plant world that these graded series of 
homonjmous parts in individual organisms are most strik- 
ingly seen. It occurs in what is perhaps its most typical, 
least modified expression in the arrangement and size rela- 
tion of parts in the leaves of many ferns and palms. But 
the compound leaves of innumerable flowering plants illus- 
trate it very beautifully. Figures 58, 59, and 60 (Acacia, 
Vicia and Cassia) show three types of compound leaves 
based on the mode of gradation of the leaflets. These might 



The Unity of the Organism 

be described as the bi-gradicnt, the direct gradient and the 
reverse gradient types, depending on whether the gradation 
is from the mid-region of the axis both ways (figure 58), from 
the proximal toward the distal end (figure 59), or from the 
distal toward the proximal end (figure 60). 



Almost all simple leaves of seed plants show something of 
the same scheme. As examples, typical elliptical-entire 
leaves of the elm and poplar and such typical lobed leaves as 
those of most oaks may be pointed to. 

Nearly every twig of a tree which represents a single 
annual growth impulse, in cases where the growing period is 
restricted to a small part of each year, presents a size gra- 
dient in the leaves distributed along the axis. A particularly 
striking illustration of this is furnished by the California 

Growth Integration 


coast redwood, Sequoia semperrirens (figure 61), where the 
new segment is short, is added end on to the one before it 
until a considerable succession of segments is produced, and 
where the leaves are retained for several years. That each 
segment in these leaves is an annual production is not cer- 
tain, probably several segments being sometimes formed in a 


single season ; but however that may be, that growth occurs 
in a series of impulses, each of which is sharply recorded 
in the size gradations of the repeated parts, is obvious 

It is a familiar fact, too, that in many plants a similar 
quantitative gradation of the reproductive parts along an 
axis occurs, but the extent to which this scheme pervades 
the constituents of the members of the series, even to the 
seeds, appears not to have attracted much interest on the 
part of botanists. To illustrate this point I present a single 
set of measurements, one of many which I have collected, 

The Unity of the Organism 

of parts of the fructiferous organs of plants. These meas- 
urements are of Frasera perryi, an abundant annual in 
southern California and rather specially favorable for such 
a study in that the fruit stalk is single in each plant, stands 
up intact and rigid after it is fully ripe and dry, and is al- 
most mathematically regular in the disposition of its parts. 
The table was compiled from measurements of a single plant, 
and three measurements pertaining to each seed vessel are 
given, namely, the length of the interval on the main axis 
between each two vessels, the length of the pedicels which 
bear the vessels, and the length of the vessels themselves. 
The measurements are all in millimeters. Several other di- 
mensions might have been taken, which would almost cer- 
tainly have produced similar results. 


Length of 

Length of 

Length of 

























































15 15 21.5 13 

That these gradations hold, at least in some plants, even 
to the seeds is certain as the following tabulation of the 
weight of seeds from different parts of the seeding axis of a 
wild mustard plant (Brassica nigra) shows. The figures 
were compiled from the weights of seeds taken from groups 

Growth Integration 103 

of twenty capsules from the bases, middle portions, and dis- 
tal ends, respectiA r ely, of six such stalks. The weights given 
are in grams. 

Total weight Number of Seeds Av. Wt. per Seed 

Base .903 950 .00095 

Middle .694 ?'>'! .00088 

Tip .330 525 .00063 

This mere glance at an exceedingly common phenomenon 
in living nature must suffice for the present. 

Justification for Bringing All These Phenomena Under One 


Probably about the first question that most persons would 
raise concerning what we have presented would be as to how 
far the series dealt with have anything to do with one an- 
other. Especially, we may apprehend, would most biologists 
question the justifiability of bringing together the raeristic 
phenomena in animals and the repetition of parts in plants. 
If such a collocation of phenomena must be justified on the 
basis of known causal factors, then undoubtedly is justifica- 
tion impossible in the present state of knowledge. But justi- 
fication of this sort is not called for by the point now oc- 
cupying us. What concerns us at present is the quite for- 
mal fact that when any lot of homonymous objects fall into 
a quantitatively graded series the members of that series 
have a fixed relation to the series as a whole. They are not 
interchangeable with one another. Each is a function, 
mathematically speaking, of its set or series. Vertebra m 
of the python's skeleton, myotome m of the amphioxus body, 
tube-foot in of the starfish arm, branchlet m of the ascidian 
tentacle, leaflet m of the vetch leaf or of the redwood shoot, 
seed-vessel m of Frasera, seed-lot m of the mustard plant, 
and m, or any other member you choose from any other 

104 The Unity of the Organism 

series whatsoever, is a determinate thing; it is what it is 
partly because of its position in the series regardless of 
whether the physical or other producing agent of the dif- 
ferent series be the same or wholly different. 

Even this purely structural formal basis establishes 
the fact of a measure of integratedness for all individual 
organisms in which the phenomenon appears. But to leave 
the subject at that would be superficial and unsatisfactory 
indeed. However, reflection makes it almost certain that 
there is some sort of causal basis for the phenomena. This 
conclusion follows first from the fact that the series result 
from the growth of the organism, and second from the cer- 
tainty, at least in many cases, that the continuance of life 
of the individual involves the maintenance of the series, this 
in turn involving some measure of metabolic interdepen- 
dence among the members of the series. 

Attempted Causal Explanation of These Series 

For establishing the general truth of this type of inte- 
gration we need not, in strictness, go any further than we 
have gone. Nevertheless, the importance of the subject 
justifies a few remarks on attempts that have been made to 
explain the series causally. The best known of these comes 
from botanists, and conceives that the diminishing series of 
leaves and other structures, seen with more or less distinct- 
ness almost universally among plants, is due to the increas- 
ing remoteness of the successive parts from the roots of the 
plant, that is, from the main source of the plant's food. It 
is obvious, however, that this explanation is not of general 
application, since in animals the food does not come from a 
root system which anchors the organism to its food-yielding 
medium. Nor is it possible to bring the series in all animals 
into correlation with a blood circulatory system, as their 
existence in many coelenterates, hydroids and alcyonarians 

Growth Integration 105 

for, example, where no circulation exists, shows. It is al- 
most certain, too, that the series occur in many plants that 
have no sap system such as is assumed by the physiological 
explanation above indicated. Many of the marine algae 
come under this head, a striking example of which is the kelp 
Macrocystis pyrifera of the western coast of both Americas. 
That the laminae of this plant fall into a beautiful direct 
gradient series is a fact which can not escape the notice 
of any one who sees them. The question of whether each 
streamer of laminae reaches finally and necessarily a limit of 
growth in which the size series is present is not so certain, 
but from considerable attention to the question I am almost 
sure this is the case, although the point needs more study. 
Another interesting and probably useful course of reason- 
ing about organic growth attempts to connect the results of 
growth with autocatalytic chemical action. Although these 
attempts have not, so far as I am aware, taken special cog- 
nizance of the natural size series which are occupying us, 
but have been concerned with the weights or volumes of or- 
ganisms at various stages of growth, there is little doubt 
that the phenomena we have been considering are closely 
connected with those dealt with in these attempts. T. Brails- 
ford Robertson seems to have devoted more thought to this 
matter than any one else. The following, taken from the 
summary of conclusions found in his original paper, pre- 
sents the most essential parts of his theory: "(1) In any 
particular cycle of growth of an organism or of a particular 
tissue or organ of an organism the maximum increase in vol- 
ume or in weight in a unit of time occurs when the total 
growth due to the cycle is half completed. (2) Any particu- 


lar cycle of growth obeys the formula log K(t-ti), 

A x 

where x is the amount (in weight or volume) of growth 
which has been attained in time t, A is the total amount of 
growth attained during the cycle, K is a constant, and t^ is 

106 The Unity of the Organism 

the time at which growth is half completed." 

Assuming that the growth of an amphioxus, let us say, to 
adulthood represents a growth cycle of this statement, that 
the production of somites begins with the most anterior one 
and proceeds toward the tail, and that the successive growth- 
increments (corresponding to x in the formula) are regis- 
tered in the somites as we find them, then the animal's body 
as exhibited by its myomercs would correspond fairly well 
to Robertson's statement under (1), as would several of the 
other growth series we have glanced at, and as would also 
great numbers of series presented by ordinary plants and 

The formula for growth contained in (2) is, according 
to Robertson, u such as would be expected to hold good were 
growth the expression of an autocatalytic chemical reac- 
tion." Assuming the general correctness of these state- 
ments, no one interested in the larger problems of organic 
growth could hesitate to believe that they must be impor- 
tant in some way. 

However, that the relations shown do not prove that au- 
tocatalytic chemical activity is a cause of growth in any- 
thing more than a subordinate, contributory way, is obvious 
on reflection. In the first place, Robertson himself has 
pointed out, in substance, that such action says nothing 
about the particular shape which the mass of transformed 
substance takes, but since some characteristic configuration 
or shape is fundamental to all organic growth, the entities 
for which A and x stand in the formula are really only ab- 
stractions. Although the formula may apply approximately 
to a great many organisms, it will apply to none exactly, 
except by chance to a very occasional one. This is the gen- 
eral form of criticism, illustrations of which are seen in the 
fact that in the series of direct and inverse gradients shown 
in the vetch (figure 59) and Cassia (figure 60), respectively, 
the formula appears not to apply at all. The general pur- 

Growth Integration 107 

port of the strictures here placed upon the value of this 
explanation of growth is well brought out by Moeser, who 
says, probably with literal truthfulness, "One will not find 
two germinating plants (Keimpflanzen) which would have 
exactly the same growth curve, even though they proceed 
from seed of absolutely the same weight and grow under 

*/ o o 

exactly the came conditions." 4 

The kernel of this criticism is that even though it should 
be established, as very likely it will be, that autocatalytic 
action is an essential factor in all growth, it can be a causal 
explanation in only a partial and subordinate sense. This is 
so because although K is a constant for a particular indi- 
vidual as observed, it assumes different values for different 
groups, partly at least because no account is taken of size 
or configuration. Moreover, even if these factors were con- 
sidered, there would be nothing corresponding to a physical 
constant (depending only on autocatalytic action), since 
one of the most distinctive things about organic growth is 
that it is differential, the differentials corresponding to the 
taxonomic group to which the individuals belong. 

Even though we are still much in the dark as to what the 
causal nexus is between the growth processes and the quan- 
titatively graded series so widely seen in nature, it seems 
certain that some such nexus exists, and that its operation 
constitutes a true integrational factor in the individual. It 
appears, too, that this type of integration is about the 
simplest and that it accompanies the simplest type of dif- 
ferentiation, the two together constituting the simplest type 
of organization above the organization of the cell. But into 
this interesting subject we can not now go. 

Axial Metabolic Gradients as Integrative Phenomena 

For the most thorough and sustained experimental inves- 
tigation of the primary integrative processes in growing 

108 The Unity of the Organism 

organisms that has been made, biology is indebted to C. M. 
Child. In two recent volumes he has summed up and sys- 
tematized the elaborate researches prosecuted by him in this 
field almost exclusively for fifteen years, and has presented 
his conclusions more fully than in any of his previous writ- 

The limitation set for the present to the constructive part 
of our enterprise makes it impossible to do more than touch 
at a few points the great mass of experimental evidence on 
which Doctor Child bases his conclusions. Fortunately, 
however, the kernel of the conclusions can be stated rather 
clearly in a small space. 

Although (surprisingly, it seems to me) Child refers 
hardly at all to the graded meristic series occurring in na- 
ture, to which the preceding pages have been devoted, it 
can hardly be doubted that the phenomena with which he 
deals, and calls "axial gradients," come under the same head 
as do those which have been occupying us. The phenomena 
which in the first instance Child has been concerned with, 
have been brought to light mainly through studies on re- 
generation in many lower animals. But the general con- 
clusions reached are far broader than this ; indeed they ex- 
tend to well-nigh the whole scope of organic growth, but 
especially to growth which involves elongation either of the 
whole organism or of certain parts of organisms. Thus the 
head-tail type of individual, whether the body be segmented 
as in arthropods and in many worms, or unsegmented as in 
other worms and in molluscs, is perhaps the most striking 
exemplification of the axial gradations with which Child 
deals. The following quotation shows the generality with 
which he views the matter from the ontogenic side: "Gra- 
dients in rate of cell division, size of cells, condition or 
amount of protoplasm in the cells, rate of growth, and rate 
and sequence of differentiation are very characteristic fea- 
tures of both animal and plant development. Such gra- 

Growth Integration 109 

dients are definitely related to the axes of the individual or 
its parts, and are evidently expressions of axial metabolic 
gradients. While the existence of such gradients indicates 
the existence of gradients in activity of some sort, the 
various kinds of gradients are not all necessarily pres- 
ent where metabolic gradients exist. In some cases the vis- 
ible gradient may be a gradient in rate of growth or in 
protoplasmic constitution ; in still others a gradient in 
sequence of differentiation, etc., and sometimes metabolic 
gradients exist without any structural indications of their 
presence. At best these various kinds of gradients are 
merely general indications of differences in metabolic rate 
and undoubtedly in many cases the visible differences along 
an axis represent something more than differences in meta- 
bolic rate. The important point is that visible indications 
of graded differences in metabolic rate occur so generally 
in definite relations to the chief axes of the bodv. ;> 

* ' 

One phase of this general statement is the developmental 
correlation that various regions of the bodv in many lower 

^j . *s 

animals have with the head or anterior end, these regions 
being developmentallv dominated, in Child's expression, by 
the anterior end proportionally to the distance of the re- 
gion from the end. 

A typical case is furnished by flat-worms of the genus 
Planaria, animals especially favorable for experiments in 
regeneration, since they are very hardy to laboratory con- 
ditions and have great powers of reconstituting themselves 
from pieces of various sizes, shapes and positions cut from 
them. "Any piece of the planarian body," says Child, "is 
capable of giving rise to all parts posterior to its own level, 
whether a head is present or not, but no piece is capable of 
producing any part characteristic of more anterior levels 
than itself, unless a head begins to form first." 

From a great mass of experimental evidence produced by 
Child and others we have the following: "These facts force 

110 The Unity of the Organism 

us to the conclusion that in such experimental reproductions 
there is a relation of dominance and subordination of parts. 
The apical or head region develops independently of other 
parts but controls or dominates their development, and in 
general any level of the body dominates more posterior or 
basal levels and is dominated by more anterior or apical 
levels." 7 

A really unique merit in Child's work is the fact that he 
has given special attention to the connection of these axial 
gradients manifesting themselves in various structural and 
functional ways, with the fundamental metabolism of the 
organism. Several methods of experimenting have been em- 
ployed to this end, the one most frequently used being what 
he calls the susceptibility or survival-time method. The es- 
sence of this depends upon the fact, determined by many ob- 
servers, "that a relation exists between the general meta- 
bolic condition of organisms, or their parts, and their sus- 
ceptibility to a very large number of substances which act 
as poisons, i.e., which in one way or another make meta- 
bolism impossible, and that difference in susceptibility may 
be used with certain precautions and within certain limits as 
a means of distinguishing differences in metabolic condition, 
and more specifically, differences in metabolic rate." 

The demonstration of metabolic gradients by this method 
depends upon the fact that "death and disintegration of dif- 
ferent parts of the body usually follow a regular sequence," 
this making it possible "to determine the time, not merely of 
disintegration of the whole animal, but of the various re- 
gions of the body." 

Another way of showing difference in rate of metabolism 
in different parts of the organism is by the use of the biom- 
eter, an apparatus for estimating minute quantities of carbon 
dioxide, recently devised by S. Tashiro in connection with 
his important researches on carbon dioxide production in 
nerves. By these methods it is shown, pointing to a single 

Growth Integration 111 

instance, that in pieces of a flat-worm isolated by cutting 
"the rate of metabolism is higher in long anterior pieces 
than in posterior pieces of the same length." 

Starting from this low but seemingly universal level of 
integrative phenomena in the individual, Child formulates 
views of the nature of organisms that agree very well with 
the organismal standpoint upheld in this volume. He 
writes : "The organic individual appears to be a unity of 
some sort. Its individuality consists primarily in this unity, 
and the process of individuation is the progress of integra- 
tion of a mere aggregation into such a unity, for this unity 
is not simply the unity of a chance aggregation, but one of 
a very particular kind and highly constant character for 
each kind of individual. In all except the simplest individ- 
uals it determines a remarkable degree of uniformity and 
consistency, both in the special relations of parts and the 
order of their appearance in time, and also in coordination 
or harmony of functional relation to these parts after their 

M eristic Gradients and Metabolic Gradients Both 
Phenomena of Growth Integration 

In view, then, of the exceedingly wide prevalence in living 
nature of axially disposed meristic series quantitively 
graded, and of the equally wide or even wider prevalence of 
axial gradients on the basis of metabolic activity, the 
gradients of both sorts arising as fundamental growth phe- 
nomena, it appears impossible to avoid recognizing our first 
category of integration, namely, growth integration, as 
about the most simple and primal and universal of all these 
categories, at least for multicellular organisms. It seems 
as though the other kinds of differentiation and integration 
are superposed, as one might express it, upon this primordial 
kind. To a consideration of the other, superimposed inte- 

The Unity of the Organism 

grations we now pass, taking them again in their seeming 
order of obviousness. 


1. Torrey 139 7. Child (1) 215 

2. Ritter ('09) 71 8. Child (1) 66 

S.Robertson 612 9. Child (1) 77 

4. Moeser 373 10. Child (1) 73 

5. Child (2) 65 11. Child (2) 2 

6. Child (1) 213 

Chapter XVIII 


Functional as Contrasted with Growth Integration 

A SSUMING Child's theory of metabolic gradients to be 
* *-well grounded, we are furnished thereby with one im- 
portant insight into the chemical processes involved in the 
unity of the individual organism. But that process is con- 
cerned primarily with the growth, with the production of the 
individual. The question now is, are there chemical proc- 
esses the object of which is to maintain the functional unity 
of the complete or nearly complete individual? Are there 
chemical operations the office of which is to preserve a 
proper interrelation among the parts of the organism as 
these perform their special offices? 

That such is to some extent the significance of most if 
not all internal secretions as usuallv understood is indicated 


by the fact that the functional disturbances attending re- 
moval of the thyroids or other glands from various animals ; 
and by the further fact that where internal secretions play 
a part in development, their action is rather that of stimula- 
tor, or at least modifier, than of true producer. 

The conception of internal secretions as being at least as 
much regulators of physiological function as of growth is 
illustrated by cases of hypopituitarism of the post- 
adolescent type, like those described by Gushing, for ex- 
ample. In the series of cases of disease due to "pituitary 
deficiency" the first symptoms appeared when the subjects 
were from thirty to forty years old. 


114 The Unity of the Organism 

The Conception of "Internal Secretions" 

The nature of the phenomena now to be considered, and 
their significance for our discussion make it desirable to 
think about these secretions from the broad standpoint first 
stated, according to Bayliss, by Brown-Sequard and d'Arson- 
val, namely, as materials produced by any living cells or 
tissues which are discharged into the blood or lymph and 
have specific effects on other parts or functions of the or- 
ganism. Regarded thus it is now known that many cells of 
the organism produce internal secretions. Although we are 
more concerned with function than with structure in this 
discussion, our purpose will be best served by beginning 
with a morphological classification of the secretion-pro- 
ducing cells. They may be divided into tAVo categories, 
those which are disposed into definite organs, or glands, the 
ductless glands of long standing in anatomy ; and those 
which are not assembled in such organs. Knowledge of this 
second class of cells is of recent date,, and is fuller from the 
functional than from the structural standpoint. The chief 
glands, to which the name Endocrine has lately been given 
by Schafer, are now so well known as hardly to need men- 
tion. They are the thyroid apparatus, including the thy- 
roids and the parathyroids, the suprarenal body, the pitui- 
tary body, and probably the thymus and pineal bodies. Cells 
now known to produce internal secretions but which are not 
arranged in glands are certain cells of the pancreas scat- 
tered among the pancreatic cells proper ; certain cells of 
the alimentary mucous membrane; the interstitial cells of 
the ovary and of the tcstis, and probably certain cells of 
the placenta, of the mammary gland, and of the uterus. 

Following our usual course of making the treatment 
merely illustrative rather than aiming at exhaustiveness, 
our selection will include one example from each of these 
groups. From the glandular category we take the thyroid 

Chemico-Functional Integration 115 

apparatus, and from the non-glandular a portion of the 
alimentary mucous membrane, namely, that of the duo- 

Effects of Removing tlie Human Thyroid for Curative 


As definite knowledge of the great physiological impor- 
tance of internal secretions begins with human surgery- 
with operations on the thyroid apparatus we may well 
begin our study here. This is the better starting point 
in that there is no more striking illustration of how great 
a part of the whole organism may be implicated in the 
action of internal secretions than is afforded by the prod- 
ucts of the thyroids and parathyroids. 

The subject first came into clear light in the early eighties 
of the last century through the experiences of Swiss sur- 
geons, Theodor Kocher and J. L. Reverdin especially, who 
removed the thyroids to cure goitre, this disease being spe- 
cially prevalent in some parts of Switzerland. The patients 
operated on were found to improve rapidly for a time after 
the operation, but later untoward symptoms began to mani- 
fest themselves. Because the variety and pervasiveness of 
these symptoms in a typical case are highly instructive for 
us we present them in detail, selecting a description from 
Human Physiology by Luciani : "Patients who have under- 
gone total thyroidectomy . . . experience the initial symp- 
toms of glandular deficiency either at once or at latest some 
weeks after the operation. They feel weak, complain of 
heaviness of the limbs, and more or less diffuse dull pains, 
particularly in the legs, which may become acute and assume 
the character of pains in the bones. 

"Other more serious symptoms are gradually associated 
with the preceding. After four or five months the face 
and the extremities swell and become cold, the muscles are 

116 The Unity of the Organism 

torpid, sometimes rigid, often exhibiting muscular tremors, 
and are incapable of carrying out any delicate manual acts 
of precision. At first the swelling is variable ; it is more 
pronounced in the morning than in the evening, but steadily 
increases until it becomes permanent. It is not ordinary 
cedema, in which percussion with the fingers leaves a depres- 
sion ; it Js a hard and elastic swelling. It is specially local- 
ized in the hands, feet and face, where it produces a char- 
acteristic alteration of the countenance. The lower eyelids 
are the first to present a sacculated, semi-transparent swell- 
ing, which is hard to the touch ; then the infiltration spreads 
to the folds of the face, w r hich become smoothed out ; to the 
nose, which gets rounded ; to the lips which swell, and 
bulge outward, saliva dribbling from them. The features 
are coarsened and expressionless like those of a cretin. 

"The mental functions accord with this appearance, since 
they are blunted, so that the patients lose their memory, 
become deaf, taciturn, melancholy, self-absorbed, and reply 
extremely slowly to questions. They further complain of 
slight but perpetual headache ; feel an almost constant sen- 
sation of cold, which is most acute at the extremities ; at 
times they are seized with vertigo, and may even lose con- 

"All the symptoms become still further aggravated. The 
whole body may grow more bulky from the extension of the 
swelling. The skin loses its elasticity, can only be picked 
up in large folds, and becomes dry owing to defective 
capacity for sweating. The epidermis desquamates in more 
or less extensive lamellae, particularly on the hands and feet ; 
the hair turns grey, falls out, and gets constantly thinner. 

"The heart functions weakly, but with ordinary rhythm ; 
the pulse is small and thready. Examination of the blood 
shows nothing constant ; but there is often a more or less 
pronounced and progressive oligocythaemia, which undoubt- 
edly contributes to the characteristic pallor of the skin, 

Chemico-Functional Integration 117 

this being of the earthy, yellow-spotted hue peculiar to 

"The respiratory rhythm is almost always normal ; the 
digestive apparatus functions well, as also the urinary 
system. The spleen is not enlarged." 

This complex (syndrome in medical terminology) of 
manifestations is known technically as cacliexia tliyreo- 

Later experience by other surgeons with the same opera- 
tion discovered that in some cases the effects are much more 
acute and rapid, and may be replaced by what has been 
called "tetany" (though having little in common with ordi- 
nary tetanus), ending in death more often than otherwise. 

Experimental Thyroid Excision in Normal Lower Animals 

No sooner had the far-reaching influence of the thyroid 
for the human organism begun to be recognized in this way 
than experimentation on inferior mammals was invoked for 
further light on the subject. Moritz Schiff, the ground- 
breaker in this field, published in 188-t the results of the 
removal of the thyroid from a large number of dogs. In 
all cases where the whole thyroid apparatus was excised 
the dogs soon died after a run of such symptoms as tremor, 
spasms, and convulsions. Nor did Schiff rest content with 
merely ascertaining the effects of removal, complete and 
partial, of the thyroid apparatus. He found that these 
effects could be entirely prevented by grafting a portion of 
the gland under the skin or into the body cavity of the 
animal before the thyroid operation, or by injecting thyroid 
juice into the blood or lymphatic systems, or by feeding raw 
thyroid to the dogs. The story of how these experiments 
led to the now widely practiced treatment of myxoedema with 
thyroid or thyroid extract would be out of place here, 
though it should not be passed wholly unnoticed. 

118 The Unity of the Organism 

So an enormous mass of evidence, experimental, surgical 
and clinical, is now in court demonstrating 1 that for some 


animals at least, among them being the human and the 
canine species, products of the thyroid apparatus are in- 
dispensable to the normal life, the symmetrical growth and 
balanced physiological activities of the organism. That 
the apparatus is essential to the "Hormonic Equilibrium" 
of the organism in some animals is beyond question. 

While no pretense can be made at an exhaustive exami- 
nation of this evidence two phases of our discussion make 
it desirable to cary the examination on the manifestational 
side a little farther. One of these is the importance of 
making as objective and emphatic as possible the extent of 
the manifestations in the individual ; the other is the ques- 
tion of the generality, taxonomically speaking, of the thy- 
roid apparatus. 

In the interest of the first of these I present, verbatim, the 
report of a single case of complete thyroidectomy, the animal in 
this instance being a fox. The individual concerned was a female 
less than one year old. 

"Oct. 28. Glands removed; good recovery. 

"Oct. 29- Normal., but does not eat. 

"Oct. 30. Salivation, rapid breathing, strong tremors and tet- 
any from 7 A. M. to 2P.M.; quiescent but weak during the rest 
of the afternoon. 

"Nov. 1. Normal, but rather weak; eats; no sign of tremors 
or salivation during the day. 

"Nov. 2. Restless; slight tremors; dyspnoea; does not eat. 

"Nov. 3. Some depression, but no tremors or salivation until 
4 P. M. ; does not eat. At 4 P. M. spasms appeared and con- 
tinued unabated as long as observed (7 P. M.). 

"Nov. 4. Found dead at 4- A. M. Post-mortem examination 
revealed no parathyroids nor accessory thyroids." 

As to taxonomic range and character of manifestation of thy- 
roid influence, much diversity might have been anticipated on 
general natural history grounds. As far as investigations have 
gone they realize these anticipations. A summary of results will 
serve our purpose, and this is at hand in Schafer's volume already 

Chemico-Functional Integration 119 


Concerning the effects of removal of the thyroid appa- 
ratus he says : 

"The most acute symptoms are exhibited by carnivora such as 
dogs, cats,, foxes, and wolves (Vincent), and the young of herbi- 
vora (v. Eiselsberg, Sutherland Simpson) and are of a nervous 
nature. . . . Some species exhibit no symptoms whatever at 
least when the operation is performed on the adult. Horsley 
states that this is the case with birds and rabbits; but according 
to Gley, the latter are affected if care is taken to find and re- 
move all four parathyroids, and Doyon and Jouty obtained typical 
tetany in hens which had been parathyroidectomized. . . . Ac- 
cording to Vincent and Jolly badgers are totally unaffected by 
complete removal of both thyroids and parathyroids." 

From the anatomical characteristics of the organs, 
and from the known effectiveness of minute portions of 
them, such statements as the last must be taken with re- 
serve. Although these results show by their diversity that 
an enormous amount of study remains to be done on the 
comparative side, they leave no question that the secretion 
of the thyroid apparatus is important for the general 
health and equilibrium of most animals in which it occurs. 
The measure of this importance in the eyes of some authori- 
ties is seen in such a statement as, "No cell anywhere in the 
body can reach morphological perfection without thyroid 
stimulus." 4 

The Internal Secretion of the Duodenal Mucous Membrane 

We now pass to an examination of the effects of the 
internal secretion of the duodenal mucous membrane. This 
particular secretion is selected for the reasons that it is, 
according to Bayliss, one of its discoverers, "the most 
typical of all the chemical messengers"; that it was one of 
the first to be investigated ; and that it is one of the few 
which have been isolated as definite substances. 

120 The Unity of the Organism 

The mode of operation of this secretion is tersely stated 
by Bayliss : "Food entering the duodenum causes the pro- 
duction of a special substance which enters the blood and 
excites the pancreas to pour into the duodenum a digestive 
juice." 5 

That the presence of various substances, especially acids, 
in the duodenum, induces a flow of pancreatic juice was 
known when Bayliss and Starling began their work in this 
field ; but up to that time the excitation of the pancreas to 
such action was supposed to be through a nerve reflex. 
These investigators had reported in 1902 6 that acid in the 
duodenum is able to cause the pancreas to secrete after 
nervous communication between the intestinal wall and the 
pancreas is excluded. They went further and obtained an 
extract from the duodenal mucous membrane which, being 
injected into a vein, induced a copious flow of pancreatic 
juice. The substance, whatever it is, which acts thus they 
call secretm. It has been surmised by a few physiologists 
that the effects are not due to the direct action of the secre- 
tion on the pancreatic gland-cells, but that the influence 
is exerted through the vaso-dilator mechanism. But this 
surmise is negatived by the demonstration that the secretin 
will induce the flow of pancreatic juice while it does not 
alter the blood pressure. The case seems, then, fully estab- 
lished, and is so clear-cut and relatively simple an instance 
of the coordinated functioning of two wholly distinct parts 
of the body through chemical means, that it is desirable 
to get sharply before us the known steps in the process. 
In the course of normal digestion, food acidulated in the 
stomach passes into the duodenum. Here, probably in 
virtue of its acidity, it acts upon the cells of the mucous 
membrane in such a way as to induce them to produce a 
substance which is discharged, not into the intestines, there 
to take its part in digestion, but into the blood. By the 
blood stream the substance is carried through its whole 

Chemico-Functional Integration 

circuit, hence through the lungs and so on, around to the 
pancreas, the typical gland-cells of which it excites into 
activity, so that the pancreatic juice, an "external" instead 
of an "internal" secretion, is poured into the duodenum to 
exercise its digestive office on the same food which started 
the cycle of activities. 

It was with this substance particularly before their minds 
that the authors adopted the name hormone to designate 
substances which act thus. "The group of substances re- 
ferred to," says Bayliss, "which includes adrenaline and the 
various internal secretions, is characterized by the prop- 
erty of serving as chemical messengers, by which the activity 
of certain organs is coordinated with that of others. They 
enable a chemical correlation of the functions of the organ- 
ism to be brought about through the blood, side by side 
with that which is the function of the nervous system." 7 

This reference to the side-by-side activities of chemical 
messengers and nervous system in integrating the organism 
touches a subject of the utmost importance. Considera- 
tion of it must, however, be deferred until we have looked 
a little more into the nature of hormones. 

The Nature of the Active Substances in Internal Secretions 

That the peculiar iodine-rich albuminous substance ob- 
tained from the thvroid by Baumann in 1895 and since 

*< / 

observed by other investigators, contributes in some essen- 
tial way to the action of the secretion of this gland is the 
belief of apparently a large majority of authors (Bayliss, 
Eppinger, Howell, etc.), but not of all (Luciani). In view 
of the uncertainty on the point Schafer's proposal "to ex- 
press our ignorance by a term which implies no theory" 
may well be accepted, with the proviso that the term pro- 
posed be really taken as evidence that something though 
not everything is known about the substance. A part of 

The Unity of the Organism 

the proposal is worth quoting. "I propose therefore pro- 
visionally to apply the word thyrine to denote the active 
principle, whether it be identical with or contained in the 
iodothyrin of Baumann or not." 

Thyrine then becomes the name of a substance the source 
and some of the activities of which are known, but whose 
main physical and chemical attributes are unknown. 

Concerning the mode of action of thyrine there are sev- 
eral divergent views, all based on some evidence and so 
perhaps not entirely antagonistic. Is the antitoxic theory 
of Luciani partly right, right as regards the parathyroid 
secretion (Moussu, Vassale and Generali), and partly 
wrong, wrong as to the secretion of the thyroid proper, this 
being trophic rather than antitoxic? ] May there not be 
more in the enzymic theory suggested some years ago, than 
later writings have been inclined to favor? Does the fact 
that internal secretions seem to be simpler than enzymes, 
as indicated by their greater resistance to heat, preclude the 
possibility that their normal mode of action is of the enzyme 
type after all? That is, may it not be necessary to extend 
the conception of enzymic action (which is surely generic 
anyway) to include the various sorts of activity presented 
by hormones, understood in the sense given it by its origi- 

But neither can the resemblance of internal secretions 
to drugs, so far as their action is concerned, be overlooked. 
This has been dwelt upon by Schafer 5 and has important 
bearings on the problems of the origin as well as on the 
chemical nature of the substances. 

Another aspect of the mode of action of internal secre- 
tions is that of whether the effects are to stimulate or inhibit 
the activity of the organ or tissue on which they operate. 
Schafer and others make a special point of this, directing 
the attention to the fact, by way of illustration, that the 
adrenaline of the suprarenal medulla causes contraction of 

Chemico-Functional Integration 

the plain muscle of the blood vessels and inhibition of that 
of the intestines. 12 The distinction has an undoubted natu- 
ral grounding, and so is in the interest of accurate descrip- 
tion and clear conception. 

As to the actual chemical composition of internal secre- 
tions, knowledge is exceedingly meager. More is known 
about adrenaline, the active principle of the suprarenal 
gland, than about that of the secretion of any other gland 
or tissue. This was isolated by the Japanese chemist 
Jokichi Takamine in 1901, and has since been more fully 
examined by several investigators, notably by T. B. Aldrich. 
It is described as a micro-crystalline substance occurring in 
at least five crystal forms. Aldrich assigns to it the em- 
pirical formula C 9 H ir ,NO ;] , this structure placing it not far 
from tyrosin in the benzene or aromatic series. Of special 
interest is the astonishingly minute quantites which produce 
physiological effects. According to Aldrich 0.000001 gram 
of an aqueous solution of the chloride per kilo of body 
weight injected into the blood system raises the blood pres- 
sure 14 mm. of mercury. 1; 

The chemical nature of Tetlielin, lately isolated from the 
anterior lobe of the pituitary of the ox, has been studied 
by its discoverer, Robertson. It is described as white or 
pale cream colored, readily powdered, highly deliquescent, 
and having a greasy odor and slightly acid reaction in 
aqueous solutions. It contains 1.4 per cent of phosphorus 
and four atoms of nitrogen for every atom of phosphorus. 
The phosphorus-nitrogen content of the substance is con- 
sidered by Robertson as specially significant, since this 
seems to ally it chemically with "phytin," a substance found 
in the rapidly growing parts of plants, and in milk. The 
natural suggestion is that the growth-promoting substances 
in plants, milk, and the pituitary secretion are chemically 

The Unity of the Organism 

The Close Resemblances and Interrelations of the Different 

Internal Secretions 

Even the meager, merely illustrative examination of in- 
ternal secretions we have been able to make brings out the 
close resemblance there is between the several endocrine 
glands, and also between the physiological effects of the 
various secretions. These resemblances suggest an intimate 
organic interrelationship among all the internal secretion- 
producing parts of the body. 

All investigators in this field, no matter to how restricted 
a section of it their efforts are primarily directed, seem to 
come upon the interdependence of the sources and activities 
of hormones. To illustrate, Gushing, whose central interest 
has been the hyphophysis, is led to conclude that experi- 
mentally induced hyphophyseal deficiency works histologi- 
cal changes in many if not all the other ductless glands. 
It is not surprising, consequently that far-reaching theories 
have been elaborated on the basis of these relationships. 

Bayliss refers with approval to Elliott for the conserva- 
tism with which he sums up the present state of knowledge on 
this aspect of the general subject. But even so, features are 
pointed out "which suggest a common bond" : 

"(1) Carbohydrate metabolism is influenced, not only by 
the pancreas, but also by the thyroid in super-activity, in 
acromegaly, and by the injection of adrenaline. 

"(2) Growth is affected by the testis and the cortex of 
the suprarenals, arrested by the absence of the thyroid. 

"(3) Nervous implications. 

"(4) The pituitary becomes hypertrophied when the thy- 
roid is removed. Acromegaly may lead to enlargement of 
the thyroid." 14 

At the other extreme of what may be regarded as legiti- 
mate scientific theorizing, we have the views of Sajous, who 
believes research will finally demonstrate a relationship be- 

Chemico-Functional Integration 

tween all the ductless glands the combined functioning of 
which dominates most of the activities, normal and patho- 
logical, of the organism. 

Sajous' elaborately worked-out theories of internal se- 
cretions, especially in their relation to disease and medical 
practice, are opposed at many points to prevailing opinion 
based on present day research. Nevertheless regarded from 
the standpoint of general biology there would seem to be 
much merit in his effort on the one hand to find a common 
ground in the metabolic processes for all the phenomena at- 
tributed to endocrinal activity ; and on the other hand to 
find a more consistent morphological and physiological basis 
of definition and classification of internal secretions and the 
structures which produce them than has yet been recognized. 
For example, whether Sajous is right or not in contending 
that the pituitary body does not produce an internal secre- 
tion, certain it is that the non-glandular structure of its 
posterior part, extracts of which alone have slowing effects 
on the heart, is strongly suggestive to the critical naturalist 
that the inclusion without qualification of this part at least 
of the organ among the endocrine glands is an instance of 
what is known to taxonomists as "lumping" in classification 
-a kind of practice that advance in knowledge always finds 
to be inadequate for purposes that are critical. Sajous' 
late summary of his views is highly suggestive to the general 
biologist, even though it is excessively theoretical in some 

As far as a much interested outsider can judge, the pres- 
ent state of understanding of the relationships among the 
internal secretions is set forth with exceptional judiciousness 
by Waller. "There can be little doubt," the author opens 
his discussion by remarking, "that the various internal secre- 
tions are most closely correlated, yet perhaps the most dif- 
ficult, and also the most fascinating problem of present day 
medicine, is to assign to each its proper and right share of 

126 The Unity of the Organism 

importance." 15 This statement, coupled with the fact that 
one of the main objects of the discussion is to display the 
many contradictions which the author's large experience as 
a practitioner has found in the action of the internal se- 
cretions, is about the most striking, and from our stand- 
point most significant thing about this paper. 

As one illustration of the agreement of action of the se- 
cretions, or at least of the influence of the endocrine glands, 
it is pointed out that changes in calcium metabolism have 
been observed after removal of the thymus ; in disease of the 
pituitary and of the pineal bodies ; after castration ; after 
ovariotomy ; after removal of the suprarenals, and after re- 
moval of the thyroids and parathyroids. 

Of the numerous instances of contradiction which he 
brings out, we mention only that concerning tetany. This 
he shows may result from either removal of the thyroid or 
from an overdose of thyroid extract. The explanation of 
the contradictions in the action of a given secretion favored 
by the author is that of the "varying influence of the other 
internal secretions." But the descriptions given seem to 
leave no doubt that difference in type of individuals also 
comes into the explanation. Thus among children afflicted 
with enlarged tonsils and adenoids, the two distinct types 
dependent upon the character of the symptoms, is a case in 
point. One type is dull and stupid, stunted in growth, has 
dry coarse skin, and may display symptoms of rickets. The 
other type is vivacious physically and mentally, given to 
peevishness, irritability and quick fatigue, and always want- 
ing a change of activity. This type is over-tall for its age, 
perspires readily and is fine-skinned. Both types of cases 
are benefited at least for a time, Waller says, by treatment 
with thyroid extract. 

Concerning the general nature of the interdependence 
among internal secretions, this author's views seem to me so 
eminently sound that I cannot refrain from quoting them in 

Chemico-Functional Integration 

some fullness : "Considerable stress has been laid upon the 
antagonism of different internal secretions by various au- 
thors. I believe we should gain a truer insight into their 
working if we dwelt rather upon their harmony. It does not 
strike me as a very high conception of the human organism 
that health should consist in the balance of dissentient or 
antagonistic forces. It would seem far more ideal that all 
the internal secretions should work together for the common 
good of the organism, and that when some special demand is 
made upon a particular gland the others will work in har- 
mony with it. Every gland is probably necessary for the 
perfect activity of the rest, and the harmony between the 
glands is demonstrated by physiological experiments. . . . 
When it is found that the removal of an organ constantly 
induces either atrophy or hypertrophy of some other organ, 
we can reasonably deduce that in the first case the organ 
removed is essential to the welfare of the one that atrophies 
in its absence, and in the second case that the hypertrophying 
organ is endeavoring to replace the lost one, in some de- 
gree, and that therefore the two organs have a kindred 
function." 16 

As an example of the first case, the fact is cited that the 
removal of the thyroid or of the anterior part of the pitui- 
tary induces the atrophy of the testicles or the ovaries. The 
second case is illustrated by the hypertrophy of the supra- 
renal from the removal of the thyroids, and also by the 
hypertrophy of either the thyroid or the pituitary on re- 
moval of the other. "The demonstrated facts of hyper- 
trophy," we read, "clearly point to an entente or even a 
triple alliance between thyroid, hypophysis and suprarenals. 
And the genital system is absolutely dependent upon the 
integrity of these three." 1T 

Stating now, in a single paragraph, the results of inves- 
tigations in this field, we have : The different parts and ac- 
tivities of the organism are maintained in their normal state, 

128 The Unity of the Organism 

both as to the essential nature of each, considered individ- 
ually, and as to their relation with one another, by a number 
of exceedingly powerful and subtle chemical substances (in- 
ternal secretions) which are produced by certain parts, are 
passed into the blood, and by it are carried about over the 
whole organism to exert their appropriate influences on 
other parts and functions. Because of the peculiar way these 
substances do their work, they have been called chemical 
messengers, or to have a distinctive name, hormones. 

Relation Between the Internal Secretory and Nervous 


But no physiological truth is better known than that one 
of the main offices of the nervous system is to correlate the 
organs and parts of the body with one another. It is but 
natural to suppose, therefore, that if there is a chemical 
scheme for accomplishing the same end, the two are in some 
way closely related. 

That the relations which exist between the cerebro-spinal 
nervous system, the autonomic nervous system (including the 
sympathetic), and the internal secretions, constitute one of 
the most important subjects in the whole physiological do- 
main, at the same time that it is one of the most recondite 
and difficult to investigate, has come gradually to view 
through the work of the last few decades. We will try to 
extract enough from the mass that has been written on the 
subject, to illustrate the principles involved. The modern 
period of knowledge of what was formerly but rather in- 
definitely included under the term sympathetic nervous sys- 
tem, has revealed that we have to do with a portion of the 
general nervous mechanism which in reality is a subdivision 
of a larger category. 

Chemico-Functional Integration 129 

Composition and Nature of tlie Autonomic System 

The name autonomic was given to this category by Lang- 
ley. "The autonomic nervous system," he says, "means the 
nervous system of the glands and of the involuntary muscles ; 
it governs the 'organic' functions of the body" ; and further : 
"The word implies a certain degree of independent action, 
but exercised under the control of a higher power." Bay- 
liss adds: "It is necessary to be quite clear that the au- 
tonomic system includes the sympathetic, since some writers 
abroad use the name as applying to all the visceral nervous 
system other than the sympathetic, speaking of sympathetic 
and autonomic." 

Perhaps the most important single fact which differen- 
tiates the autonomic from the cerebro-spinal system is the 
intercalation, everywhere in the autonomic system, of an 
extra neurone between the cerebro-spinal nerve and the part 
innervated. Cannon states this distinction very clearly: 
"The skeletal muscles receive their nerve supply direct from 
the central nervous system, i. e., the nerve fibers distributed 
to these muscles are parts of the neurones whose cell-bodies 
lie within the brain or spinal cord. The glands and smooth 
muscles of the viscera, on the contrary, are, so far as is now 
known, never innervated directly from the central nervous 


system. The neurones reaching out from the brain or spinal 
cord never come into immediate relation with the gland or 
smooth muscle cells ; there are always interposed between the 
cerebrospinal neurones and the visceral extra neurones 
whose bodies lie wholly outside the central nervous 
system." 20 

Cannon's suggestion that these interposed neurones may 
function as transformers for impulses received from the cere- 
brospinal system should be noted here. 

Three sharp subdivisions of the autonomic nervous sys- 
tem are recognizable. One is known as the vagal or cranial 

130 The Unity of the Organism 

autonomic, because it is largely made up of fibers from the 
vagus, or tenth pair of cranial nerves. Another is the sym- 
pathetic, or better, the thoracico-lumbar autonomic, be- 
cause its fibers originate from the great visceral sympathetic 
ganglia. This is by far the most extensive of the three sub- 
divisions, and is the only one that is distributed to all parts 
of the body. The third is the sacral autonomic. As its name 
implies, it is quite restricted in extent, its fibers being dis- 
tributed to the extreme distal end of the intestine, the urin- 
ary bladder, and some of the external genital organs. But 
the differences between the three which are most important 
for us are physiological, a particularly important difference 
being that the thoracico-lumbar division acts antagonistic- 
ally to both the end divisions. Stimulation of the fibers of 
the sympathetic has just the opposite effect to the same 
stimulus applied to the fibers of the others. "The sympa- 
thetic fibers check, the vagal autonomic fibers excite, the 
movements of the intestines ; the sympathetic dilates, the 
vagal autonomic contracts, the pupil ; the sympathetic 
hastens, the vagal autonomic slows, the heart." 21 The 
sacral contracts the lower part of the large intestine and re- 
laxes the outlet of the bladder, while the sympathetic relaxes 
the same part of the intestine and contracts the same part 
of the bladder. Cannon states the general principle thus : 
"When the mid-part meets either end part in any viscus 
their effects are antagonistic" 

While the incompleteness of knowledge in this field needs 
emphasizing, yet that knowledge is sufficient to put some of 
the main features beyond question, and to make clear the 
great importance of the subject and of fuller knowledge on 
it. Touching these general aspects Professor L. F. Barker 
writes : "While we do not yet understand the exact mechan- 
isms of association among the activities of the cerebrum, 
the endocrine glands, and the reciprocally antagonistic au- 
tonomic domains and their end-organs, we can begin to see 

Chemico-Functional Integration 131 

the paths which must be followed in order that more exact 
knowledge may be gained." 

Experimental Evidence of Connection Between the Adrenal 
Glands and the Nervous System 

Some of the most important information we have in this 
field is furnished by Cannon and his collaborators concern- 
ing the secretion of the adrenals and its relation to the 
autonomic and central nervous systems. It had been proved 
before Cannon began his investigations that adrenin injected 
into the blood has exactly the same effect on certain parts 
of the organism as does the sympathetic autonomic nerves 
with which the same parts are supplied, and that the effect 
of the secretion is direct and not through the nerves. In 
other words, it had been proved that the organism has two 
methods by which the same activity of certain of its parts 
can be induced, one nervous, the other chemical. Thus the 
dilation of the pupils, the erection of hairs, the inhibition 
of activities of the alimentary canal, and the liberation of 
sugar from the liver can be induced either through sym- 
pathetic autonomic centers or by the secretion of the adrenal 
bodies. This in itself was important evidence of interrela- 
tion between the nervous system and internal secretory sys- 
tem. But the experimental researches prosecuted in Can- 
non's laboratory have proved that a connection exists be- 
tween the autonomic-adrenal phenomena and the cerebro- 
spinal system through the sensory nerves, and with the 
psychic life of the animal ; and have shown the probable sig- 
nificance of the entire scheme for the life of the organism as 
a whole. 

To be a little more specific, they have proved : 
(1) That strong excitation of sensory nerves stimulates 
reflexly the adrenal glands and causes them to pour an 
increased amount of adrenin into the blood. 

132 The Unity of the Organism 

That emotional excitement (as the fright of a cat by 
a dog) similarly increases the flow of adrenin. 

(3) That this increase of adrenin in the blood may in- 
crease the liberation of sugar from the liver into the blood 
to such an extent as to make sugar appear in the urine, thus 
demonstrating a true "emotional glycosuria." 

(4) That the increased adrenin of the blood thus pro- 
duced is probably advantageous to the organism in that it 
enhances its ability to meet special stresses that naturally 
accompany special excitement, as of fear, anger, or pain, 
this advantage consisting partly in augmentation of the 
working energy of the muscles, probably through the sugar 
delivered to them, and in increasing the coagulability of the 
blood, thereby reducing the danger from bleeding wounds. 

Summing up his conclusions as to utility, Cannon writes : 
"These changes in the body are, each one of them, directly 
serviceable in making the organism more efficient in the 
struggle which fear or rage or pain may involve; for fear 
and rage are organic preparations for action, and pain is 
the most powerful known stimulus to supreme exertion. The 
organism which with the aid of increased adrenal secretion 
can best muster its energies, can best call forth sugar to 
supply the laboring muscles, can best lessen fatigue, and can 
best send blood to the parts essential in the run or the fight 
for life, is most likely to survive." 24 

But fear and rage are, one hardly need be reminded, in 
part psychic phenomena, and hence inseparably connected 
with the higher centers of the cerebrospinal nervous system. 
Though in the main reflex and automatic, they are neverthe- 
less to some extent subject in man to intelligent control. 
Thus the way is open for a measure of rational understand- 
ing of the structural-functional means by which human be- 
ings "tap," as William James would say, and bring under 
direction those remarkable "reservoirs" of ordinarily unused 
energy about which everybody know r s something from his 

Cliemico-Functional Integration 133 

own experience, and upon which nobody has written more 
intelligently than James. 

An excellent beginning has been made, then, in the ex- 
perimental demonstration of the integration . of the endo- 
crinal and common glandular systems, the blood circulatory 
system, the autonomic and cerebrospinal nervous systems, 
and the emotional-psychic life of animals. 

Clinical Evidence of Adrenal-nervous Connection 

But important knowledge and general views in this field 
are also coming from clinical medicine, and pharmacology. 
A general presentation of the results reached down to 
1913 is contained in Innere Secretion und Nervensystem, 
by H. Eppinger and others. A particularly significant body 
of evidence coming from this source concerns the relation 
between the sympathetic or middle autonomic nervous ap- 
paratus and the two end autonomic systems, the cranial and 
sacral. These two groups act, it will be recalled, antagon- 
istically to each other. Eppinger and others have shown 
that the thoracico-lumbar, or sympathetic, and cranial, or 
vagal systems differ in susceptibility to stimuli in different 
individuals, and perhaps in the same individual at different 
times, thus making the two groups what is called sympar 
tlieticotonic and vagotonic with reference to each other, de- 
pending on whether the sympathetic or the vagal is the more 
susceptible to stimuli. This difference can be demonstrated 
by the administration of various drugs, as adrenin and pilo- 
carpin. But it is known, according to Eppinger, that the 
thyroid toxin stimulates both the sympathetic and the vagal. 
From this it results that over-stimulus of either may occur 
through this source, and go to the extent of producing the 
characteristic symptoms of Basedow's or Graves's disease 
(rapid heart beat, exophthalmia, diarrhea, etc.). These 
symptoms may occur in varying degree, depending on 

134 The Unity of the Organism 

whether the patient is sympathetico- or vagotonic, the sym- 
patheticotonic type of the disease being characterized by 
marked protrusion of the eyeballs, especially rapid heart 
beat, absence of sweats, diarrhea, and disturbance of the 
respiration; while the vagotonic type is characterized by 
slight protrusion of the eyes and increase of heart action, 
by outbreaks of sweat, diarrhea, and by faultiness in the 
respiratory rhythm. 

While some observers, like Falta, do not believe the facts 
now known can be definitely classed in this manner, the ef- 
fort, justified by some positive knowledge, has at least the 
merit of specifying to some extent the intricate reciprocal 
action between the thyroid apparatus and the nervous sys- 
tem, and also between the different portions of the auto- 
nomic system; and to this extent all students of the subject 
bear witness. Thus Falta : "In my opinion everything speaks 
for the fact that in Basedow's disease the entire nerr>ons 
system is in a condition of over-excitement and that the 
pictures presented by the vegetative nervous system are as 
uncommonly manifold and always changing." 

The indication of prime importance in this is that in these 
antagonistic divisions of the autonomic nervous and endo- 
crinal glandular systems, operating together with the other 
portions of the organism, there is a balancing-off or equil- 
ibrating apparatus through which the whole complex of vege- 
tative functions is carried on, all of which in turn are con- 
nected with the psychic functions. Probably no better il- 
lustration can be found of the conception of the organism as 
fundamentally dynamic. According to this conception nor- 
mality, both in function and in structure, consists not in 
rigid, invariable activities and organs, but in a ceaseless play 
of constitutively antagonistic forces and structures. By this 
conception the w r hole life of the organism, physical and psy- 
chical, may be crudely likened to the performance of the 
tight-rope walker, which depends on numberless balancing 

Chemico-Functional Integration 135 

activities. Let the performer be really motionless in every 
part for one instant, and he falls. 

The treatment of tetany and its relation to internal se- 
cretions, especially to that of the thyroid, by Eduard Phleps 
in the work now under consideration, is another excellent il- 
lustration of how interpretation may run in accordance with 
this conception of the animal organism. The essence of the 
section, as touching this question, is contained in the fol- 
lowing : 

"On the ground of clinical symptoms authoritative clin- 
icians like Eulenburg, Kahler and Nothnagel explained tet- 
anus as a disease of the entire nervous system. Later it was 
proved that this disease was not due to primary organic 
changes of the nervous system, but to secondary functional 
disorders. . . . The view of those authors, who refer the 
disease to the simple effect of a substance of the epithelial 
granules acting normally and continuously on the whole 
nervous system, finds here a further development of that old 
theory because for them the clinical picture of a regular 
grouping of nervous stimulus- and response-phenomena 
arises from an impairment of the close functional relation 
between the nervous system and the epithelial granules (Mac- 
Callum, Chvostek jun., Biedl, Eppinger, Fait a, Rudinger, 
Jonas, et al.). In agreement with these authors we con- 
ceive the action of the epithelio-secretive substance as that of 
a hormone in the sense of Starling and Bayliss, which must 
have its essential point of attack on certain reflex stations 
of the central nervous system." 

Taking cognizance, now, of the fact that most if not all 
the cells known to produce internal secretions arise embry- 
onically from epithelium, as does also almost all nervous 
tissue, we have the suggestion of a deep-seated combination 
scheme, chemical-and-nervous, for integrating the organism. 
The best investigated example of what is here referred to is 
the suprarenal bodies. It is now fully established that the 

136 The Unity of the Organism 

inner or medullary portion of the organ, the part which pro- 
duces the adrenin, is developed from the same neuroblastic 
mass out of which the sympathetic autonomic ganglia arise. 
Furthermore, it seems beyond question that the so-called 
chromophil, or sometimes the adrenin granules in the chief 
cells of the functioning gland play a fundamental role. Com- 
bining these facts with the equally well ascertained facts 
that the cortex of the gland is derived from the same em- 
bryonic mesoblastic mass which gives rise to the genital 
glands ; and that in adults cortical changes in the supra- 
renals are intimately correlated with reproductive changes, 
and a general view of the factual basis on which the sugges- 
tion rests is before us. We may, I think, regard the sugges- 
tion not only as justified but as revolutionary in comparison 
with any theory that was scientifically justifiable until re- 
cently. The following further quotation from Phleps brings 
the idea into still clearer view : 

"The unqualified dependence of the nervous system on the 
epithelial bodies (this last used in the sense of general physi- 
ological considerations, and only by way of illustration), 
and the absolutely vital significance of this enables us to see 
many things in a new light. We learn that many motor, 
sensory and vasomotor-trophic functions of the central nerv- 
ous system even up to the highest reflex stations having the 
most complicated cortical functions, are in constant func- 
tional cooperation with organs which heretofore have not re- 
ceived sufficient consideration from this standpoint. The 
results compel a change of our views concerning the con- 
stantly dominating position of the nervous system. We may 
see in many of its activities only the most manifold inter- 
mediary roles between glandular functions which are in the 
closest relation to metabolism, and the sum total of all re- 
actions which follow external stimuli." 27 

Chemico-Functional Integration 137 

Summary of Present State of Knowledge In This Field 

Speaking generally, we may say that the trend of all re- 
sults, experimental and clinical, is unquestionably toward a 
demonstration of the closest interaction between the entire 
internal secretory system and both the autonomic and cere- 
brospinal nervous systems, this interaction affecting the 
whole of both the growth and the functioning of the animal 

But we must remind ourselves again how fragmentary 
knowledge is in this great realm. Unanswered questions meet 
one on the very threshold of any portion he enters. By 
way of illustration, take the phenomenon of abnormal growth 
known as acromegaly. This malady is characterized, as the 
name indicates, by a "peculiar non-congenital hypertrophy 
of the upper and lower extremities and of the head." 2S Such, 
according to Schafer, is the definition given by Pierre Marie, 
who first fully described the disease. The main visible symp- 
toms consist in the enlargement of the bones of the head, 
hands, feet, chest, etc., especially in their terminal portions. 
Through such growth the nose and lower jaw, especially the 
chin, become protrudent. But the whole skeleton is more or 
less affected, and there is a corresponding over-development 
of the muscles, the affected person becoming abnormally 
strong. That acromegaly is constantly associated with an 
abnormal condition of the hypophysis is recognized by ap- 
parently all authorities. Whether the abnormality of the 

*/ +] 

gland is a cause or only an accompaniment of the disease is 
an open question in the minds of some. However, the view 
of a large majority is that such a causal relation does exist. 
"That the acromegalic skeletal growth," says Schafer, "is 
produced by hypertrophy and oversecretion (or perverted 
secretion) of the anterior lobe is highly probable, both as the 
result of partial extirpation in animals and from the effect of 
operative removal of the pituitary tumours in man." 29 The 

138 The Unity of the Organism 

particular issue here, it will be noticed, is the vitally im- 
portant one of what might be called the functional as 
distinguished from the hereditary cause or at least incitement 
of growth. 

In illustration of the importance of understanding the 
unification among these complex systems, the manifestation 
of which is in turn dependent upon the organism as a whole, 
the following from the address by L. F. Barker already re- 
ferred to, is impressive. 

"In how far these sudden and violent excitations of the 
autonomic nervous system which accompany strong emo- 
tions are due to the intervention of the glands of internal 
secretion, and in how far they depend upon direct neural 
conduction from the brain, we are as yet but ill-informed. I 
need only remind you of the vasodilation of the face in the 
blush of shame, of the palpitation of the heart in joy, of the 
stimulation of the sudoriparous glands which precedes the 
sweat of anxiety, of the stimulation of the vasoconstrictors, 
the pupil dilators and the pilomotors in the pallor, mydriasis 
and goose-skin of fright, to illustrate some of these violent 
autonomic excitations." 30 

The references here, it will be noted, are primarily mani- 
festations pertaining to the surface, the integumentary 
parts of the body, and their scope is what especially inter- 
ests us. Very nearly the whole list of these parts is in- 
volved, and probably a complete inventory would be still 
more inclusive. 

Now notice the range of manifestations at a deeper level 
that are involved. "The balance maintained normally be- 
tween the two antagonistic systems the vagal and sympa- 
thetic autonomic is one of the most interesting of physio- 
logical phenomena. Think, for example, of the rate of the 
heart-beat how constantly it is maintained at a given level 
in each individual when the body is at rest ; the impulses ar- 
riving through the vagal system just balance those arriving 

Chemico-Functional Integration 139 

through the sympathetic system, so as to maintain a rate of 
approximately seventy-two beats per minute. And a similar 
balance is maintained in other autonomic domains (e.g., the 
pupils, bronchial musculature, gastric glands, gastro-intes- 
tinal muscle, sweat glands, bladder muscles, etc.)." 30 

And Barker then calls attention to the extent to which the 
normal processes of the body depend upon temporary upsets 
of these equilibria, examples of which are watering of the 
mouth at the smell or sight of food which appeals to the 
appetite through these senses, the flow of gastric and pan- 
creatic juices at the proper time, through indirect stimula- 
tion ; the sudden relaxation of the sphincter and contraction 
of the detrusor of the bladder in micturition ; the violent 
contractions of all the muscles concerned in parturition in 
the female, and so on. 

We may summarize the results of this chapter thus : 

(1) The researches of recent years on the internal secre- 
tory system and its connection with the great subdivisions 
of the nervous system, and with the blood, muscular, and 
reproductive systems, have laid a solid foundation for an un- 
derstanding of the chemico-ftmctional basis of the animal 
organism's unity. 

(2) The emotional phase of the psychic life of the ani- 
mal is proved to be in direct organic connection with this 

From these results there naturally springs the important 
question : What relation has human consciousness to this 
same basis? An attempt to answer this question will be an 
unavoidable part of our treatment later of the psychic in- 
tegration of the organism. 


1. Luciani II, 13 5. Bayliss 707 

2. Carlson and Woelfel 49 6. Bayliss and Starling .... 325 

3. Schafer (1914) 18 7. Bayliss 706 

4. Hertoghe 194 8. Schafer 28 


The Unity of the Organism 

9. Luciani II, 22 

10. Luciani II, 35 

11. Schafer 9 

12. Schafer 11 

13. Aldrich 457 

14. Bayliss 721 

15. Waller 277 

16. Waller 280 

17. Waller 281 

18. Langley 240 

19. Bayliss 484 

20. Cannon, W. B.('16) 22 

21. Garrison 144 

22. Cannon, W. B. ('16) ... 34 

23. Garrison ; 146 

24. Cannon, W. B. ('13) ... 372 

25. Falta 59 

26. Eppinger, et al 214 

27. Eppinger, et al 217 

28. Schafer 106 

29. Schafer 69 

30. Garrison 146 

Chapter XIX 


T remains now to consider the real purpose for which 
the presentation of facts and views on internal secre- 
tions has been made, namely that of showing critically the 
significance of this secretory system for the organismal 

General Inability of Elementalism to Interpret the 


There is perhaps no better way of approaching this part 
of our task than by noticing the inability of elementalistic 
biology to deal in a really intelligent and consistent manner 
with the phenomena in this field. The breakdown of bio-ele- 
mentalism when confronted with the phenomena of "chemical 
messengers" nowhere finds more cogent illustration than in 
the effort to identify internal secretions with the organ- 
forming substances hypothesized by Sachs and others. 

Although in what follows the exposure of inconsistenc} T 
and fallacy will have to be drastic and may seem personal, 
the truth is it is wholly impersonal in spirit and is directed 
at a system of bad reasoning born of what might be called 
a juvenile metaphysics of the living world. 

The objective achievements of Jacques Loeb and others 
of the school he represents, in experimental biology, merit 
the admiration of all lovers of observational truth. One 
may be, too, more tolerant of their faults as reasoners 


The Unity of the Organism 

than he could be but for his recognition of their high service 
against all the traditional forms of supernaturalism. The 
real case against the school is, as I see it, two-fold. First, in 
their zeal to substitute naturalism for supernaturalism they 
fail to notice that supernaturalism is in its very essence 
finalistic, and they are led to imagine they have attained, 
or can attain, natural explanations that fully supplant the 
old supernatural explanations. This results in the con- 
version of their supposed naturalism into something which 
is essentially another kind of supernaturalism. The second 
part of the case against the school is its abuse of the most 
common principles of the knowledge-getting processes in ob- 
jective biology. For the general good of the biological 
sciences the urgent need of reformation touching both as- 
pects of the case has led me to examine the particular in- 
stance of Loeb's treatment of internal secretions at greater 
length than would otherwise be justifiable. 

Critique of the View That Internal Secretions are "Formative 

Stuffs' 9 

Although Loeb is the only author, so far as I know, who 
has expressly contended for the identity of internal secre- 
tions with Sachs' organ-forming substances, the assumption 
is so accordant with the spirit of elementalism, and Loeb is 
so typical and eminent a protagonist of this philosophy, 
that his proposal will probably find many adherents. It is 
consequently desirable to see what there is in the effort to 
bring hormones into such a historical setting. 

The statement of Loeb's views is contained in "The Or- 
ganism as a Whole from a Physico-Chemical Viewpoint," 
1916. Referring to his espousal twenty-five years ago of 
Sachs' hypothesis to explain heteromorphosis, he writes : "At 
that time the idea of the existence of such organ-forming 
substances was received with some scepticism, but since then 

Significance of the Internal Secretory System 143 

so many proofs for their existence have been obtained that 
the idea is no longer questioned. Such substances are known 
now under the name of 'internal secretions' or 'hormones'; 
their connection with the theory of Sachs was forgotten with 
the introduction of the new nomenclature." 

A case to which Loeb makes special reference as proof 
that internal secretions are the same as Sachs' organ-form- 
ing substances is that of the effect of thyroid on the meta- 
morphosis of the tadpoles of frogs and toads, demonstrated 
by Gudernatsch. The author's mode of using this case in 
illustration of his contention is highly instructive. He re- 
fers to the legless condition of the tadpoles and calls atten- 
tion to so much of Gudernatsch's results as pertain to these 
members. Gudernatsch found, Loeb points out, that where- 
as in the usual course of things the tadpoles live from four 
months to a year before the legs grow out, by feeding them 
on thyroid gland these members can be induced to appear 
at any time. "We must, therefore, draw the conclusion," 
Loeb says, "that the normal outgrowth of legs in a tadpole 
is due to the presence in the body of substances similar to 
the thyroid in their action (it may possibly be thyroid sub- 
stance) which are either formed in the body or taken up in 
the food." 2 When the case is presented in this way and 
nothing more said about it, it certainly looks considerably 
as if the thyroid substance or something like it is leg-form- 
ing substance; and such an interpretation would be enticing 
were it really true, as Loeb says, that no other substance 
seems to have such an effect. 

When, however, one consults the account given by Guder- 
natsch himself as to what his experiments were and what 
they established, the whole matter stands in a quite different 
light. First of all, the fact that in these experiments mam- 
malian thyroid was largely used as food for the tadpoles, thus 
bringing it to pass that if thyroid substance was specific 
organ-forming substance, then frogs' legs were produced by 

144 The Unity of the Organism 

mammalian substance, shows at once that there is something 
badly askew in the theory. So we are incited to examine 
the facts, and particularly the reasonings, carefully. 

The following is taken from one of Gudernatsch's papers : 
'The most striking and at the same time unquestionable re- 
sults were attained by thyroid feeding. . . . The influence 
of the thyroid food was such that it stopped any further 
growth but on the contrary led to an abnormal diminution 
of the size in the animals treated, while simultaneously it ac- 
celerated the differentiation of the body immensely and 
brought it to a premature end." 3 In other words, the effect 
of thyroid food was to stop the increase in size of the frog's 
larva and start, almost at once, its transformation into the 
adult. Now this transformation does not consist merely in 
the production of legs, but in a whole series of changes, 
some of which, like leg transformation, are progressive, 
while others are regressive. For example, one of the re- 
gressive changes to which Gudernatsch gives particular 
attention is the resorption of the tail. "Reduction of the 
body mass (resorption of the tail, loss of water, therefore 
an increasing compactness of the body, etc.)" 4 more than 
in normal development, the author says, goes hand in hand 
with the progressive changes. That is, when the entire series 
of results of thyroid feeding are considered, and not merely 
one result picked out arbitrarily, then in case we choose to 
say the thyroid substance is organ-forming as regards legs, 
we should have to say it is organ-destroying as regards 
tail. Furthermore, if we call the thyroid substance organ- 
forming, consistency would compel us to recognize that one 
and the same substance is not only formative of legs but 
of numerous other organs and parts, as of the skin, mouth, 
respiratory and blood systems, all of which undergo, as is 
well known, progressive changes during metamorphosis. Nor 
is this complex series of morphological changes the while 
story. Striking and characteristic changes in the habits 

Significance of the Internal Secretory System 145 

of the tadpoles resulted. "Towards the end of metamor- 
phosis the animals hardly moved about in the water. They 
were always lying quietly, generally on their backs. When 
disturbed they would move for a few seconds in a somewhat 
convulsive manner and then drop again to the bottom of 
the dish, while tadpoles fed on other material would swim 
about for a long time." 5 So mammalian thyroid substance 
is not only organ-forming for a whole series of frog organs 
but it is habit-forming for a variety of frog habits ! 

But we must not let the ludicrousness of this veer us 
away from the reasoning in the case. Taking the facts 
actually brought out by Gudernatsch, what becomes of the 
specificity of the substance, which according to Loeb's state- 
ments w r as what Sachs hypothesized? "Sachs suggested that 
there must be in eacli organism as many specific organ-form- 
ing substances as there are organs in the body." The truth 
appears to be that thyroid substance in this case is organ- 
forming in much the same sense that water is organ-forming 
for the leaves, flowers, and fruit in a squash vine, which could 
not develop without water. Indeed the analogy suggested 
goes further than appears at first sight. As everybody 
knows, the effect on young plants of a scant water supply 
is to stunt the plant as to size and to hasten its blossoming 
and fruiting. That is, an under-supply of w r ater has an 
effect on immature plants similar to that of an over-supply 
of thyroid substance on immature frogs, namely that of re- 
tarding growth and hastening metamorphosis. The total 
effect in each case is systemic. In other words, the real 
significance of the instance used by Loeb is just the opposite 
of his interpretation of it. Thyroid substance is organ- 
forming only through being organism- trans forming. Full 
justification of this way of interpreting the part in 
development played by thyroid substance is furnished by the 
recent studies of B. M. Allen. This investigator lias, like 
Gudernatsch, experimented with frog larvae. He has, how- 


146 The Unity of the Organism 

ever, supplemented Gudernatsch's work by depriving the 
larvae of thyroid altogether instead of giving them an extra 
allowance through feeding. Allen extirpates the entire em- 
brvonic thyroid from tadpoles long before any indications of 
metamorphosis appear. What he finds of special impor- 
tance for the present discussion is contained in the follow- 
ing : 

"While Gudernatsch showed that thyroid feeding acceler- 
ates development, this work shows that the total absence of 
the thyroid gland produces complete cessation of somatic 
differentiation at a certain stage but does not hinder con- 
tinued growth in size." 7 


The first part of this statement taken alone might be 
looked upon as confirmatory of the "formative stuff" theory 
of thyroid substance. But the phrase "at a certain stage" 
implies, as Allen's paper as a whole conclusively shows, that 
the exact opposite of such a conception is alone tenable. 
What actually happens, Allen brings to light, is that in 
spite of the complete absence of thyroid substance trans- 
formation of the larva is begun but is not carried through. 
That is, the organs and parts of the adult frog are laid 
down but (with the exception of the reproductive glands) 
are not completed. "It is evident," Allen says, "that the 
thyroid gland is in no wise essential to the earlier phases of 
development, but that at a certain definite stage, further 
development of the soma is dependent upon it." 7 It should 
be mentioned that Allen makes a rather special point of the 
accordance of his results with those of Gudernatsch. 

If the greatly hastened and modified metamorphosis of the 
frog tadpole observed by Gudernatsch is inducible by no 
other means than by thyroid substance (which while pos- 
sible is not at all certain), then is the substance causal in 
the important sense of being not only competent but in- 
dispensable. But even so it falls far short of being a com- 
plete causal explanation of the phenomena under contempla- 

Significance of the Internal Secretory System 147 

tion, as Allen's results directly prove. 

A defect in Loeb's reasoning is his ignoring the truth that 
thyroid substance lacks in the case cited the prime attribute 
of a sufficient cause, namely full competency. The actual 
substance which enters into the new legs, and as far as that 
goes, into the other new parts, is probably provided very 
little, if at all, by the specific substance or hormone of the 
thyroid. The causal role played is the relatively humble one 
of excitor or stimulator to an activity not essentially new 
but only exceptional as to time. Of course Loeb does not 
need to be told that hormones incite growth rather than 
provide the substance itself out of which the organs and 
parts are built. Indeed portions of his account of this very 
case show positively that he is aware of this. We read: 
"Thus we see that the mesenchyme cells giving rise to legs 
may lie dormant for months or a year but will grow out 
when a certain type of substances, e.g., thyroid, circulates in 
the blood. There may exist an analogy between the activ- 
ating effect of the thyroid substance and the activating 
effect of the spermatozoon or butyric acid (or other par- 
thenogenetic agencies) upon the egg." 

This suggestion of analogy between the action of thyroid 
secretion and "butyric acid (or other parthenogenetic agen- 
cies)" is well taken. The resemblance between the two 
agencies, as judged by their effects on development, is cer- 
tainly rather close. Very well; would Loeb, then, call bu- 
tyric acid organ-forming substance, and identify it with the 
"formative stuff" of Sachs ? Certainly any substance which 
will rouse the latent developmental capacities of an egg into 
activity is in a minor sense formative, especially if these 
capacities are wholly unable to start without some such 
agency. But since butyric acid, or some one of the other 
dozens or scores of parthenogenetic agencies, may activate 
the eggs of many, many species of animals ; and since the 
eggs of many, many species may be activated by any one of 

148 The Unity of the Organism 

scores of such agencies, what about the specificity of the 
formative substances which Loeb himself expressly says was 
part of Sachs' conceptions? 

Again, how reconcile the contention that thyroid sub- 
stance or something like it is organ-forming for the legs 
of a tadpole with the statement quoted a few paragraphs 
back about mesenchyme cells which give rise to legs? 

The Metaphysical and Logical Weakness of the View 

Obviously Loeb's treatment of this subject contains irre- 
conciliable contradictions. Is it then worthless? My answer 
is, no, not by any means. But how comes it that a scien- 
tist of his great experience and merited distinction can run 
into such self-destroying speculations and statements, seem- 
ingly without rational discomfort to either himself or others 
of the school which he represents? 

The answer takes us back to some of the most funda- 
mental issues between the elementahst and organismal stand- 
points, and though not requiring us to palliate in any degree 
such offenses 'against scientific reasoning, it partly explains 
how tolerance for such offenses is begotten, and discovers 
a nucleus of genuine merit in Loeb's position. Bringing the 
matter to as basal a statement as possible, what we find is 
that this whole book on the Organism as a Whole is written 
on' the theory that the only alternative to the assumption 
of supernaturalism is materialism. Instead of supernatural 
forces of some sort (Platonic Ideas, entelechies, psychoids, 
"supergenes") for explaining the organism when regarded 
as alive and whole, his assumption is that material elements 
as known to us in inorganic nature are the sufficient causal 
explanation of organic phenomena. 

Were this theory correct- -were it true that the "vital 
principle" must either be conceived as supernatural or that 
the inorganic elements taken by themselves are competent 

Significance of the Internal Secretory System 149 

to produce organisms, then it would be impossible for biol- 
ogy to do much better in its reasoning and general attitude 
than Loeb and other elementalists do when they undertake 
to construct a philosophy of organisms. I agree whole- 
heartedly that all supernaturalism, no matter what nomen- 
clatorial garb it takes on, must be repudiated by the sciences 
of organic beings. Ideas, or psychoids, or entelechies, or 
"principles" of an} 7 kind conceived as independent of, or 
even separable from, sensible objects are quite as repugnant 
to me, an organismalist, as they are to any element alist. The 
essence of my contention is that the natural substitute for 


these imponderable things are the living, individual organ- 
isms themselves, and not the particles of which they are com- 
posed. Each and every individual organism is a natural 
reality by exactly the same criteria that the atoms, mole- 
cules, cells and tissues of which it is composed are natural 
realities. And since each individual is to some extent differ- 
ent from every other, and maintains its individuality in full 
possession of these differences, by its power of transforming 
foreign substance into its own substance, it is ultimate both 
as to structure and as to causal power in as deep and literal 
a sense as the material particles of which it is composed are 

Loeb's considerable attention to the views of Claude Ber- 
nard is fortunate for us, since it affords a chance to show 
from still another angle the inevitable breakdown of ele- 
mentalist reasoning when it is brought face to face with or- 
ganic phenomena as actual nature presents them to the 
modern student. Loeb calls attention, properly, to the fact 
that one of the things on which Bernard placed special em- 
phasis, as Bichat before him had done, is the organizing syn- 
theses which go on in the living being. The real advance, it 
seems to me, which Bernard made over any of his predeces- 
sors, was the positiveness of his rejection of a vital force 
as something supernatural as something, using his own 

150 The Unity of the Organism 

words, "under the government of a special principle, a pecu- 
liar power what name soever be given it, whether soul, or 
archeon, or psyche, or plastic intermediary, or guiding 
spirit, or vital force, or vital properties" ; and his rejec- 
tion of these conceptions because of his recognizing that the 
unitariness of the organism removes the necessity for as- 
suming any such extraneous principle. "When we say," 
Bernard writes, "that life is a guiding idea, or the evolu- 
tive force of the being, we merely express the thought of a 
unity in the succession of all the morphological and chemical 
changes effected by the germ, from the beginning to the 
end of life. Our mind grasps that unity as a conception it 
cannot escape. . . ." 10 

Up to this point the position held by Bernard is entirely 
satisfactory for the organismal conception as I am trying 
to develop it, and is, according to my view, unassailable. 
But the unquoted part of the last sentence contains a state- 
ment which reveals Bernard on a by-road leading away from 
the promised land toward which he was headed as long as he 
was speaking in terms of biology proper. The rest of the 
sentence follows : "and explains it as 'a force' ; but the mis- 
take is in supposing that this metaphysical force acts after 
the manner of a physiological force." Stated in a nut- 
shell, the by-road which Bernard is entering here is that of a 
kind of separatedness, but inevitable concomitance, or paral- 
lelism between the phenomenal and neumonal worlds which, 
according to the views upheld in this volume, does not exist. 
"We need here," Bernard says, "to draw a distinction be- 
tween the metaphysical world and the phenomenal physical 
world, which serves as its basis, but which can borrow noth- 
ing from it." This Leibnitzian theory, according to which 
"everything takes place in the soul as though there were no 
body, and in the body everything takes place as though 
there were no soul," Bernard says science "recognizes and 
adopts in our day." But in our day, this year nineteen 

Significance of the Internal Secretory System 151 

hundred and eighteen, science, at least so much of it as 
speaks through this volume, though understanding fully 
what this dualist theory is, rejects it. It denies that things 
take place in the soul as though there were no body ; and 
that things take place in the body as though there were no 
soul. On the contrary it affirms that what in rather un- 
critical language we call "the body" and "the soul" are in 
the most intimate and indissoluble connection with and de- 
pendence upon each other, and together constitute "the 

The Form of Metaphysical Absolutism Involved 

A full, systematic justification of this position is beyond 
the province of this volume ; and in this chapter we are con- 
cerned solely with the "body," the strictly morphological 
and physiological aspect of the subject. Nevertheless, this 
much of contact with the "soul" aspect was unavoidable for 
the reason that Bernard and also Loeb have run into it in 
such fashion as to color deeply their discussions and out- 
look. This coloring is the more unfortunate and the more 
insistent in requiring attention from the fact that the au- 
thors, especially Loeb, are apparently unaware of such 
coloring. For example, Loeb writes on the first page of his 
book, after saying that the atomistic theory of matter and 
electricity are now in all probability on a "permanent 
basis" : "This permits us to state as an ultimate aim of the 
physical sciences the visualization of all phenomena in terms 
of groupings and displacements of ultimate particles, and 
since there is no discontinuity between the matter consti- 
tuting the living and non-living world the goal of biology 
can be expressed in the same way." 

Statements like this, many of w^hich can easily be quoted 
from Loeb's writings, leave no question about his meta- 
physical affinities. The conception of "ultimate particles" 

he Unity of the Organism 

as explanation of all phenomena, is exactly what I mean by 
elementalist absolutism. 

Confusion of Theory of Organisms and Theory of the 

Knowledge of Organisms 

Although the thoroughgoing metaphysical character of 
these statements is evidenced by the finalism which crops 
out at several points, this is not the aspect of the matter 
which chiefly interests us here. Rather what we are con- 
cerned with is the fact that affirmations about the "aim of 
the physical sciences" and the "goal of biology" do not 
belong, properly speaking, to the provinces of physical 
science and biology at all, but to quite a different science, 
namely that which deals with the nature of knowledge itself. 
The "physical sciences" are the vast accumulation of man's 
positive knowledge, theories, hypotheses, and so forth, about 
physical nature; they certainly are not physical nature it- 
self. Consequently a statement of the character and aims 
of that knowledge is not a statement about the phenomena 
to which the knowledge pertains. And the same reasoning 
applies to the affirmation about the goal of biology. 

All this is only another way of saying what Loeb him- 
self virtually tells us, namely that his entire discussion of 
the organism as a whole is made from the standpoint of one 
particular theory of the ultimate nature of living beings, 
that theory being the mechanistic. Recall the complete title 
of the book The Organism as a Whole From the Physico- 
Chemical Viewpoint. To treat the subject from this view- 
point is of course perfectly legitimate. When, however, the 
assumption is made that such a treatment is the only really 
legitimate one because it rests on ultimate truth, then sound 
science is bound to protest, chiefly because of the obvious 
biological inadequacy, and at some points, perversion and 
contradiction displayed in the treatment. 

Significance of the Internal Secretory System 153 

This brings us back to the way internal secretions are 
dealt with by Loeb. His failure to distinguish between the 
two very distinct fields of theory above indicated, namely 
theories about the phenomena of living beings, and theories 
about knowledge of these phenomena, largely explains the 
defects of the theoretical parts of his work. And we are 
now in position to give our criticism greater explicitness. 
Consider for example in the light of what has just been said 
about confusion relative to kinds of theory, the irreconcil- 
able statements, cited on an earlier page, that the mesen- 
chyme cells give rise to the legs of the tadpole, and that 
thyroid substance is organ-forming substance for the legs. 
Stated briefly the case seems to be thus : Loeb recognizes, as 
every one must, that internal secretions constitute a physico- 
chemical agency for bringing about that harmonious de- 
velopment and functioning so characteristic of the organ- 
ism. But this harmony is one of the very things which has 
seemed to some biologists inexplicable without the assump- 
tion of supernatural influences of some sort ; hence Driesch's 
attempt to modernize the ancient entelechy. But since in- 
ternal secretions play the role that entelechies are supposed 
to play, namely that of establishing and maintaining the 
unity and equilibrium of the organism, the need for ente- 
lechies no longer exists ; at least, this would be so for all 
persons who do not contend that "ultimate explanation" is 
the "goal of science." 

To those who hold these absolutist beliefs as to the power 
and aims of science three favorite courses are open and are 
followed by different representatives of the school, depend- 
ing on the taste, training and outlook of the person. One 
course consists in pointing out, taking an illustrative case, 
that internal secretions, being only contributing causes, do 
not constitute an ultimate explanation, so that entelechies 
or something similar are as necessary as before. This would 
be the course followed by the vitalistic wing of the absolu- 

154 The Unity of the Organism 

tist school. For them internal secretions would be, in so far 
as they contribute to the harmony of the organism, merely 
agencies produced and used by supernatural causes. 

Another course, and perhaps the one most frequented by 
elementalists, would be to contend that internal secretions 
are sufficient as a causal explanation of organic unity to 
make the entelechy or any similar notion quite superfluous, 
even though these substances are far from a complete ex- 
planation. The reasoning of this group of elementalists as 
to this situation is substantially as follows : although in- 
ternal secretions fall far short of fully explaining organic 
unity and harmony, the action of these being merely that 
of incitors and inhibitors, they are yet genuinely causal, 
genuinely physico-chemical and so are on the road toward 
complete explanation of the phenomena. All that is nec- 
essary consequently, is to believe that still further advance 
in the same direction will reach finallv a full element alis tic 


explanation ; that is, an explanation which will have no need 
of either supernatural elements or the organism as a whole. 
The attitude of this large class is one primarily of faith 
rather than of reliance on positive knowledge ; they are 
inspired more by what they believe they will do in the future 
than by what they actually have done. They are preemi- 
nently men of promises. Although their achievements in 
experimental science are indeed large, the results reached 
by them are prized more on account of what they are believed 
to augur for the future than for their present meaning and 

Then, finally, there is the group of elementalistic absolu- 
tists, of whom the author of Tlie Organism as a Whole from 
the Physico-Chemical Viewpoint is one of the most eminent 
in our day, who, as we have pointed out, by confusing theo- 
ries about objective phenomena with theories about the 
knowledge of such phenomena, are led to affirm that such 
phenomena as the unity of the organism are fully explained 

Significance of the Internal Secretory System 155 

by internal secretions, when as a matter of fact they are 
only partly explained thus. 

The surprising thing about the confusion of this group 
is that wholly irreconcilable positions are held with impun- 
ity, such for instance as those according to which thyroid 
substance is held to be the organ-forming substance of frogs' 
legs in one part of a discussion, and mesenchyme cells are 
acknowledged to be of this nature in another part. The 
contradiction is, to be sure, often of such character as 
easily to escape the uncritical reader; but as to the au- 
thors of such contradictions no other explanation seems 
possible than that of wrong habits of scientific thought be- 
gotten of untenable a priori conceptions. For example, a 
hasty reading of the discussion under review might lead one 
to suppose that thyroid substance is not, after all, regarded 
by Loeb as anything more than one contributing cause of 
frogs' legs, mesenchyme cells being recognized as another 
cause. Close attention to the text does not, however, war- 
rant this generous interpretation of the author's position. 
Going back to his espousal of the theory of Sachs and other 
botanists as to organ- forming substances, we read : "Specific 
shoot-producing substances are carried to the apex, while 
specific root-producing substances are carried to the base of 
a plant. When a piece is cut from a branch of willow the 
root-forming substances must continue to flow to the basal 
end of the piece, and since their further progress is blocked 
there they induce the formation of roots at the basal 
end." n 

If this means anything it means that the shoots and roots 
are actually built up by material carried about in the wil- 
low branch. There is nothing in the language that can be 
interpreted as meaning that shoot-and-root-forming sub- 
stances are mere stimulators of some other substances which 
become the actual shoots and roots. Yet it is with forma- 
tive substances of this sort that Loeb in some parts of 

156 The Unity of the Organism 

his discussion explicitly identifies internal secretions. "At 
that time the idea of the existence of such specific organ- 
forming substances was received with some skepticism. . . . 
Such substances are known now under the name 'internal 
secretions' or 'hormones' ; their connection with the theory 
of Sachs was forgotten with new nomenclature." Then 
follows the reference to the tadpole legs ; so that were consis- 
tency really a jewel to the author, he could not escape mean- 
ing that the substance of the legs was actually derived di- 
rectly from the particular internal secretion in question, in 
this case thyroid substance ; and the later statement about 
the formation of legs from dormant mesenchyme cells 
through the mere activating effect of thyroid substance is by 
implication contradicted. 

This brings up again a matter about the interpretation 
of development which we have dwelt upon in several other 
connections, that of protest against rejecting the indubi- 
table evidence of the senses in favor of a priori conceptions. 
The crucial question in the present case is this : which is the 
more fundamentally organ-forming substance for frogs' legs, 
the mesenchyme cells which "though giving rise to legs may 
lie dormant for months," or the thyroid substance which 
may stimulate these cells into premature activity? While 
Loeb's discussion does not raise this question definitely, the 
implication is unescapable that thyroid substance is for him 
the more fundamental. What else is the meaning of the 
contention that this substance is organ-forming while no- 
where do we find the mesenchyme cells so designated? Yet 
the observational evidence is that the production of legs is 
accomplished through the transformation of mesenchyme 
and other cells which in the larva are not leg-substance, but 
in the adult are leg-substance. Hence it follows that so far 
as actual observation is concerned the mesenchymatous and 
other larval substances are more entitled to be called organ- 
forming than is the thyroid substance. 

Significance of the Internal Secretory System 157 

An Illustration of Neglect of Fact By Elementalist Theory 

These reflections lead to a still deeper level of the inherent 
faultiness of elementalistic absolutism in biology, the toler- 
ance which it engenders for ignoring relevant facts ; or 
stated otherwise, of arbitrarily selecting from a great com- 
plex of facts just those which suit the argument, and dis- 
regarding all the others. For example, recall Loeb's ref- 
erence to precocious leg-production in frog tadpoles as 
though the effect of thyroid feeding stood alone rather than 
as one among a great concatenation of effects, some con- 
structive and some destructive, this complex of phenomena 
constituting the metamorphosis of the young into the adult. 
Loeb's use of Gudernatsch's results amounts to a positive 
obscuration for the reader of what these important experi- 
ments really teach. Only by the culling of facts to suit the 
argument and the use of certain words and phrases, as 
"influence," "responsible for," and so on, with equivocal 
meanings, can these results be made to support the conten- 
tion that thyroid substance is specific organ-forming sub- 
stance for frogs' legs. The patent fact is that certain 
mesenchyme and other cells of the larva are organ-forming 
for legs, and there is no straightforward way of talking 
about the causes of the transformation of a given group of 
more or less undifferentiated tadpole cells into the much 
enlarged and highly differentiated group called a leg, with- 
out recognizing the whole organism as causal of the par- 
ticular transformation. Probably no set of discoveries con- 
cerning the development of the individual has ever been 
made which so objectifies the means employed by the whole 
in producing and correlating its constituent parts as those 
on internal secretions ; and not the least significant fact is 
that these substances are themselves produced by the or- 

Even were Loeb's contention valid that thyroid substance 

158 The Unity of the Organism 

is specific organ-forming substance, the indubitable fact that 
in normal development this substance is itself a product of 
the organism's activities, throws it into a very subordinate 
place as a cause of development. The truth is, the main 
upshot of the effort to explain ontogeny on elementalistic 
principles amounts to an effort to avoid recognizing the most 
positive and definite entity in the whole situation, namely 
the organism taken alive, normal and untampered with. 

A Peculiar Element alls t Objection to the Organic Whole 

'We now come to the last point to be noticed in connec- 
tion with the element alist attempt to deal with internal se- 
cretions as related to the organism as a whole. Instancing 
the familiar way in which a particular part of a flat-worm 
will give rise to a new head after being cut away from the 
original animal, when no head would have been formed at this 
place had not the animal been cut, Loeb writes : :< How does 
the 'whole' suppress all this formative power in the part be- 
fore the latter is isolated? It almost seems as if the isola- 
tion itself were the emancipation of the part from the tyr- 
anny of the whole. The explanation of this tyranny or of 
the correlation of the parts in the whole is to be found, how- 
ever, in a different influence." 

Then follows the statement previously quoted about the 
specific organ-forming substances of Sachs and other bot- 
anists, and the assumed identity of these with internal se- 

Without raising the question concerning the evidence for 
the assumption that the production of a flat-worm's head 
as indicated is dependent upon internal secretions, let us 
consider a moment the interesting conception thrown into 
the treatment that the whole flat-worm tyrannizes over its 
parts. Why this? Is it "mere rhetoric"? We are not per- 
mitted to judge it thus, for no one has pronounced against 

Significance of the Internal Secretory System 159 

this sort of thing in science more frequently than Loeb. The 
conception that the "whole" exercises a "tyranny" over the 
parts we must accept as being seriously scientific with Loeb. 
Well then, since tyranny is "absolute power arbitrarily or 
unjustl}- administered," according to the dictionary, it is 
certainly interesting to an organismalist to find so eminent 
an elementalist acknowledging the organism as a whole to be 
truly causal relative to its parts for it is hardly conceiv- 
able that even the extreme pliability of elementalist practice 
as regards the definition of words would venture to hold the 
absolute power which constitutes tyranny to be without 
causal efficiency. Power thus potent but which could cause 
nothing, not even the destruction of the parts (for surely 
the tyranny of the whole does not destroy the parts), would 
be too queer a conception for anybody to father deliber- 

But the most interesting thing about this idea of the 
"tyranny" of the whole over the parts remains to be no- 
ticed. Tyranny is not merely absolute power; it is such 
power exercised unjustly or "in a manner contrary to law 
or justice." 

Here we come, I think, to about the last ditch of the ele- 
mentalist position. On what ground does one conceive the 
power exercised by the whole organism over its parts, to be 
contrary to law or justice. According to what legisla- 
ture or court is there a law of the parts of an organism more 
just than the law of the whole? None whatever in nature, 
it must I think, be admitted. The only ground for the ele- 
mentalist's pronouncement, that the whole acts tyrannously 
toward the parts, that it acts in a manner "contrary to law 
or justice" is in the mind of the person who makes the pro- 

The truth is and it is of great importance since its 
influence reaches far beyond the confines of scientific tech- 
nicalities any scientist, especially any biologist, who is 


The Unity of the Organism 

through and through an elementalist, is necessarily a protes- 
tant against all law except the law of elements. 

The scientific elementalist is inevitably anarchistic toward 
all the most common, most objective, structures and laws of 

His faith is in the laws of the obscure or invisible world 
and against those of the everywhere visible world. Atoms 
are more real to his mind than are lands and waters, plants 
and animals. 


1. Loeb ('16) 155 

2. Loeb ('16) 156 

3. Gudernatsch 475 

4. Gudernatsch 476 

5. Gudernatsch 476 

6. Loeb ('16) 159 

7. Allen ('18) 515 

8. Loeb ('16) 156 

9. Bernard 516 

10. Bernard 519 

11. Loeb ('16) 154 

12. Loeb ('16) 154 

Chapter XX 

Neural and Not Psychical Phenomena the Subject of This 


THE fact should be firmly fixed in mind at the outset 
that in this chapter we have nothing to do with the 
organism's consciousness and volition. We are to deal with 
the nervous system purely on the basis of its physical activi- 
ties. Whether or not consciousness or something of the es- 
sential nature of it appertains to part or all of the activities 
to be treated we do not have to decide, so far as this discus- 
sion is concerned. Our task here is to be strictly anatomical 
and physiological, and not at all psychological. 

Distinction Between Developmental and Functional 


The discussion may be opened by calling attention again 
to a matter noticed incidentally in the last chapter, namely 
the relative parts played by internal secretions and the ner- 
vous mechanism in integrating the developing organism on 
the one hand, and the functioning organism on the other. 
We saw that the role of internal secretions in preserving 
developmental equilibrium in the individual is so conspicu- 
ous as to throw the strictly physio-logical role of the sub- 
stances somewhat into the background. Nevertheless as evi- 
dence, particularly clinical evidence, has made clear, their 
part in functional equilibrium is far-reaching. 


162 Tlie Unity of the Organism 

We must now point out specifically what appeared only 
incidentally in the previous discussion, namely, that the 
integrative action of hormones and of the nervous system 
are to a considerable extent the reverse of each other as 
regards their relation to development and to adult func- 
tion. That is, while harmones are, perhaps, more impor- 
tant in development and become relatively less significant 
with the advancing age of the individual, the nervous sys- 
tem plays a minor part in integrating the developmental 
processes but becomes of supreme importance in this way 
for the functioning of the adult. Or, stating the generali- 
zation in another way, integration of the organism is ac- 
complished by chemical means chiefly, and by neural means 
little or not at all, during early life, and by neural means 
chiefly and by chemical means secondarily in later life. 

One can hardly fail to see, in a general way, the bearing 
of this on the familiar truth that the life of the individual 
among the higher animals, man especially, is successively 
vegetal, animal, emotional, and finally rational and intel- 
lectual in its dominant characteristics, with the successive 
stages of earlier and later childhood, youth, and earlier 
and later maturity. 

The Author's Indebtedness to Sherrington's Work 

My dependence upon Sherrington's work in this field will 
be so great as to make it impossible to acknowledge it at 
every point. I therefore make at the outset the general 
statement that a large part of my discussion consists of a 
re-wording and rearranging of facts and ideas contained in 
this physiologist's important book, The Integrative Action 
of the Nervous System. But while my chief reliance here 
is on Sherrington's work, the writings of Cannon and his 
collaborators have been the most important source of what 
I have to say about the autonomic nervous system. Can- 

Neural Integration 163 

non's recent book, Bodily Changes in Pain, Hunger, Fear 
and Rage, has been especially drawn upon. 

The Fundament a1ity of Cellular Integration in the Reflex 


In no department of physiology do cells keep themselves 
more persistently in the attention of the student than in the 
physiology of the nervous system. But likewise nowhere 
is the fundamental dependence of cells on other cells more 
clearly seen ; for while, speaking from the standpoint of 
general functions, cells may be looked upon as individual 
units, when it comes to the study of nerve cells as such, 
that is, as constituents of the functioning nervous system, 
the individual cell is found to be no longer the basal unit. 
Viewed thus the reflex-arc and not the individual cell is the 

The distinction thus indicated is important from the or- 
ganismal standpoint and must be considered a little more 
fully. The general functions of cells to which reference is 
made above are those common to all cells, even those of the 
tissues of fully differentiated multicellular animals. No 
matter what tissue be under consideration, muscle, gland, 
epithelial or what not, so long as it is truly living, each and 
every cell assimilates, breathes, excretes, and carries on all 
the metabolic processes. Thus far each cell is an indepen- 
dent unit in a high degree. Now while all tissue cells, using 
the term tissue in its common histological sense, have a rela- 
tional or integrational function in addition to these indi- 
vidual functions, it is in the nervous system that this inte- 
grative aspect of cell life is most positive and definable. 
The reflex-arc, as the unit of the nervous system, is itself a 
combination of three indispensable parts or elements : the 
receptor, the conductor and the effector. Typically these 
structures contain at least four cells, one for the receptor, 

164 The Unity of the Organism 

two for the conductor, and one for the effector. An illus- 
tration would be a reflex-arc consisting of a tactile cell of 
a touch organ, the two cells constituting the conducting 
path, and an effector cell in a muscle. A point of special 
interest in connection with such an arc is that the dependence 
of the parts on one another is such, in the highest develop- 
ment of the arc, that the specific action of each part is 
dependent on the specific and connected action of the other 
parts. "The optic nerve itself," to quote Sherrington, "is 
unable to enter into a heightened phase of its own specific 
activity on the application of light. Initiation of nervous 
activity by light is the exclusive (in this instance) function 
of cells in the retina, retinal receptors." 1 And of course 
without brain cells as effectors for vision the specific activi- 
ties of receptors and conductors would be impossible. Not 
only would an optic nerve fiber's conducting ability be useless 
without a retinal cell on the one hand and a brain cell on the 
other, but the very conductive act itself would not be fully 

This specific dependence, as it might be called, of the 
parts of a reflex-arc is so significant that another example 
may be profitably noticed. A striking one is afforded by 
the effects of the passage of gall stones through the gall 
ducts, instanced by Sherrington, partly on the basis of his 
own studies. The excruciating pains associated with this 
malady are due to the distention of the wall of the gall duct 
by the passage through it of the mineralized organic con- 
cretions which constitute the gall stones. The point in 
this for us is the fact that though the stimulus which pro- 
duces the pain is mechanical and acts upon the wall of the 
duct, this stimulus is so peculiar that other sorts of me- 
chanical stimuli of the same tissue, even to cutting and 
wounding, give no sense of pain. Though the duct may 
be cut without causing pain, pain may be produced by 
injecting the duct to distention with neutral fluid. "Marked 

Neural Integration 165 

reflex effects can be excited [by fluid] from the very organs 
the cutting and wounding of which remains without effect." 2 
The reflex mechanism involved is adjusted to respond only 
to a stimulus of a special kind. In this it is comparable to 
the optical mechanism referred to above. 

These illustrations show something of the general nature 
of the reflex-arc as an integrated structure. But we want 
to know something about the part contributed by the 
different constituents of the arc to this nature. The spe- 
cific office of sense organs viewed as the receptor members of 
the arcs will first engage attention. "The main function of 
the receptor is therefore to lower the threshold of excit- 
ability of the arc for one kind of stimulus, and to heighten 
it for all others." 3 This principle is so important that we 
must allow no vagueness as to its meaning. It means that 
while more than one sort of stimulus may put a particular 
sense apparatus into operation, there is one and only one 
kind, namely, that to which the sense organ is naturally 
subject, that elicits the reflex in its normal or type expres- 
sion. A sense organ may be looked upon as a group of 
receptors attuned to a special stimulus, as contrasted with 
that of the general stimuli to which an organism is always 
subject by being always in contact with its environment. 

In illustration Sherrington instances the fact that the 
threshold for the touch-sensation is lower for a mechanical 
stimulus than for an electrical stimulus. Having regard to 
the whole lot of reflex-arcs of the body, we may say that 
the different sense organs constitute mechanisms of selective 
excitability for the different stimuli, each organ being so 
adapted to its natural stimulus that it responds to this 
better than to any other. This reference to the sense or- 
gans as adaptive makes it desirable to notice the fact that, 
according to Sherrington, electricity is never an adequate 
stimulus of any sense organ because it is not a natural 
stimulus for any, since it does not enter into the natural 

166 The Unity of the Organism 

environment of the organism. This is important in view 
of the large use made of electricity in experimental work 
on nerve physiology. To what extent may accepted gen- 
eralizations concerning various reactions be influenced by 
the inadequacy of the stimuli applied? 

We glance next at the physiological character of the con- 
ducting member of the arc. It is hardly necessary to say 
that typically this part is found in the nerve cords every- 
where present in the body. The most general truth of sig- 
nificance for us is the fact that having regard to conduction 
through the arc as a whole, the process is different in many 
important respects from conduction in the nerve cord alone. 
Conduction in the arc as an integrated whole is quite dif- 
ferent from conduction in the part of the arc specially 
devoted to that office. To illustrate by a quotation : "An- 
other remarkable difference between reflex-arc conduction 
and nerve-trunk conduction is the irreversibility of direc- 
tion in the former and the reversibility of the latter." 
This is only another way of expressing the well-established 
fact that nerve impulses are incapable, in the higher organ- 
isms especially, of running in more than one direction. 
Sensory excitations can go only centripetally and motor ex- 
citations only peripherally. As Sherrington points out, this 
is part of the "law of forward direction" of the neural system. 
It is fully established that nerve impulses may run in both 
directions when a stimulus is applied at any point or a 
nerve trunk, whether the trunk be motor or sensory. Taken 
by itself this fact might encourage a somewhat careless ob- 
server to think of the nervous system as more or less hit-or- 
miss in its structure and action, it being able to work one 
way as well as another, the final result being determined by 
where the stimuli happen to be applied. As a matter of 
fact, though, when we come to consider the real unit of 
neural organization, the reflex-arc, instead of any of the 
constituents of that arc taken separately, all suggestion of 

Neural Integration 167 

haphazardness disappears. Nerve conduction in a nerve 
trunk as an isolated phenomenon has neither existence nor 
meaning for the actual organism. The conducting element 
par excellence of the real unit of nerve organization is 
differentiated with reference to the other elements of that 
unit, and normally acts only in such relation. The circum- 

. / 

stance that this element is found to have the ability to act 
somewhat differently under different but abnormal rela- 
tions signifies little or nothing, so far as one can see, for 
the normal workings of the nervous system. What it does 
show is something of the diversity and plasticity of the 
organism because of its latent abilitv to act otherwise than 
in nature it does act when conditions are imposed upon it 
which are wholly new to it. 


Although the explanation of this irreversibility of direc- 
tion appears not to be known with certainty, the suggestion 
that it may be connected with a difference in permeability 
of the synaptic membrane between cells of the reflex-arc to 
certain ions, depending upon which side of the membrane is 
presented to these ions, is plausible and indicates the re- 
sourcefulness, as one might say, of structural and functional 
method by which the organization of the living being is ac- 
complished. Should this suggestion prove to be correct, 
the question would arise, how comes it that the synaptic 
membrane is thus differential in its action? And no ade- 
quate answer would be forthcoming that did not take cogni- 
zance of the fact that the membrane assumed this differential 
mode of acting as part and parcel of the differentiation and 
integration as the reflex-arc as a whole. 

Sherrington deals with a whole series of other differences 
between nerve trunk conduction and reflex-arc conduction, 
such as the phenomena of summation of stimuli, that is, 
the adding up of excitations too slight taken singly to pro- 
duce reaction, until the aggregate brings response. Such 
are rhythmic activity in response to stimuli which are not 

168 The Unity of the Organism 

rhythmic, or at least not of the same time-intervals as the 
excitations, gradation in intensity of reaction, and so on. 
To these he devotes more than a hundred pages, all under 
the common heading, "Coordination in the Simple Reflex." 
It is unnecessary to follow the matter further, sufficient 
having been given to show something of the variety of ends, 
all definitely and specifically contributory to the needs of 
the organism as such, and all accomplished through differ- 
entiation and integration of one relatively simple organic 
structure, the reflex-arc. 

The Integration of Reflex-Arcs 

Our next task is to learn something of the combinations 
among the myriads of reflex-arcs of which the higher nerv- 
ous system is composed. The definition of the reflex-arc 
given on a preceding page presents this mechanism in its 
simplest terms, and so far as the definition indicates it would 
seem possible for it actually to exist and be operative in 
this simplicity. In fact, physiologists are accustomed to 
assume such an entity and to call it the simple reflex-arc, 
and its activity the simple reflex. But while such a concep- 
tion is convenient and helpful for didactic purposes, espe- 
cially as an aid to visualizing the earliest stage of cell inte- 
gration in the evolution of the nervous system, as a matter 
of fact, according to Sherrington, it probably never exists 
in nature. "A simple reflex is probably a pure abstract 
conception, because all parts of the nervous system are 
connected together and no part of it is probably ever 
capable of reaction without affecting and being affected by 
various other parts, and it is a system certainly never abso- 
lutely at rest." 

The factual basis for the conception that one of the most 
essential distinctions between the nervous system and other 
systems of the body is the integratedness of the former, is 

Neural Integration 169 

well expressed by Donaldson : "Isolated groups of nerve- 
cells do not occur. Indeed, a group of nerve-cells discon- 
nected from the other nerve-tissues of the body, as muscles 
or glands are disconnected, would be without physiological 
significance. It is desirable, therefore, to emphasize the 
fact that by dissection the nervous system is found to be 
connected throughout its entire extent." 

I would ask the reader to consider these statements in the 
light of the cellular conception of the organism maintained 
in this volume and especially dwelt upon in the chapters on 
the cell-theory, namely, that the true way of viewing the 
organism is not as being built up of cells in the sense of 
having been constructed by the bringing together of pre- 
viously isolated cells, as a brick house is built up of bricks ; 
but rather as being composed of cells through resolving 
itself into these as it increases in size and differentiates 
itself into its organs and functions. With such a concep- 
tion Sherrington's formulation, based on the findings of 
vast observational and experimental research on the struc- 
ture and action of the nervous system, is in perfect accord. 
If we see in the completed nervous system a complex 
mechanism developing as a unit subservient at all stages to 
the needs of the organism as a whole, the myriads of reflex- 
arcs then present themselves as final, as end-stages in the 
differentiation and not as initial states ; and the universal 
organic and functional connection with one another, affirmed 
in the quotation, would be just what we might expect. 

If, on the other hand, the integral nervous system 
were built up in a literal sense, that is, by the actual coming 
into connection with one another of previously isolated 
simple reflex-arcs, such arcs ought to be demonstrable both 
in the ontogeny of higher animals and in the adults of the 
lower metazoa. 

No one should be beguiled into the notion that the readily 
observed facts of ontogeny of the nervous system, the vari- 

170 The Unity of the Organism 

ous processes, dendrites and axones, do actually grow out 
on nerve cells and bring cells into connection with one 
another and with receptor and effector cells, and that a 
functional coordination is thus finally reached does not 
exist in any way or degree in the early stages. Proof that 
the completed stages of neural integration cannot be ac- 
complished without the production of cell-growths which 
put cells into connection with one another is a very different 
thing from proof that these cells were once wholly isolated 
from one another, and that the outgrowths which establish 
the final connections were initiated by impulses which origi- 
nated wholly within the cells ; as, by way of illustration, the 
branches of two young trees standing not far apart might 
come into contact as the trees increased in size. It is a 
matter of elementary knowledge of animal development that 
in this sense the cells of the nervous system are never iso- 
lated either from other nerve cells or cells of certain other 
parts of the organism. Until we know vastly more than 
we do know of the chemical nature of intercellular substances 
and of the chemical and physical activities which go on at 
the planes of contact between cells, nothing could be more 
gratuitous and unscientific than to assume that nerve cells 
differentiate as they would were they not in some measure 
vitally associated with one another from the very begin- 
ning. Indeed, such positive knowledge as we have tends 
strongly against such an assumption. 

Recall, for example, the fundamental part chemical mes- 
sengers play in development. And Sherrington's insistence 
on the role of intercellular substance and "surfaces of sepa- 
ration" between cells in the functioning of the adult nervous 
system is much to the point for this contention. Intercellu- 
lar as w T ell as intracellular conduction must be, he main- 
tains, expected in the reflex-arc on the basis of the cell 
theory. 7 

Neural Integration 171 

The Spreading and Compounding of Reflexes 

As a practical matter study of the integrative action of 
the nervous systems never gets away from integration among 
reflex-arcs and reflexes any more than it does from the 
cellular intergration fundamental to the reflex-arc itself. 
The general nature of the study always involves the "spread 
of reflexes over a wide range of nervous arcs." The idea 
is more fully stated in the following: "This compounding of 
reflexes with orderliness of coadjustment and of sequence 
constitutes coordination, and want of it inco-ordination. 
We may therefore in regard to coordination distinguish 
coordination of reflexes simultaneously proceeding, and co- 
ordination of reflexes successively proceeding. The main 
secret of nervous coordination lies evidently in the com- 
pounding of reflexes." For the rest, all we can do or need 
do to meet the requirements of this discussion is to get at 
the main principles as illustrated by examples of this com- 
pounding of reflexes. 

As a starting point for the discussion of this larger 
aspect of neural integration, Sherrington takes what he has 
called "the principle of the common path." Basal to this 
conception is the familiar fact that a stimulus applied at a 
single point, on the surface of the body, for example, gives 
rise to a nerve impulse which may reach a great number 
of muscles or glands. A single receptor with its neurone 
must be in communication with a great number of effectors, 
some of which are very different in kind. Looking at the 
arcs from the effector ends, as one may say, it is obvious 
that the impulses reaching innumerable effectors must come 
over a single conducting course. 

On the other hand, since a given muscle or gland may be 
reached by impulses from a great number of reflex-arcs, 
the effector and its neurone must be the "common path" 
for all these impulses, often very different in quality and 

The Unity of the Organism 

source. "While the receptive neurone forms a private path 
exclusively serving impulses of one source only, the final 
or efferent neurone is, so to say, a public path, common 
to impulses arising at any of many sources of reception." 9 

An example which illustrates the general principle of the 
common path, and several phenomena incidental to this, 
and one which has been much investigated by Sherrington, 
is that of the scratching reflex in dogs. It has long been 
known that in several land vertebrates which have the 
habit of scratching the side and back of the forward part 
of the body with the hind foot, the scratching movement 
may be elicited by appropriate stimuli applied to the area 
reached by the foot as a pure reflex ; that is, in the absence 
of any chance for impulses from the brain to reach the 
parts involved in the activities. Since the scratching move- 
ment consists in bringing the hind leg forward and upward, 
and for scratching, a rhythmic movement of the foot, the 
muscles, both flexor and extensor, of the thigh, leg and 
foot must be involved. And since the reflex can be induced 
by a stimulus applied at any point within the large recep- 
tive field (i. e., nearly the whole side and back of the body), 
impulses started from various parts of the field must pass 
through one and the same neurone in the muscles concerned. 

And here comes in a fact showing another aspect of the 
integration of reflexes in this case. A stimulus at a given 
point in the field too weak by itself to elicit the reflex may 
bring it on when acting in combination with weak stimuli 
at other points in the field. Sherrington calls reflexes 
which act together in this way allied reflexes. 

Still another kind of combination of reflexes involving 
the common path principle, even more significant than allied 
reflexes, are what are known as proprio-ceptive reflexes. 
The kernel of this class of reactions is the existence of re- 
ceptors in the deep tissues of the body, that is, not belonging 
to the surface and hence not subject to stimuli from the 

Neural Integration 173 

external world, but because of their situation subject only 
to stimuli "given by the organism itself." Specifically the 
sources of the stimuli are the muscles, joints, blood vessels 
and so on, which by the regular activities are always in 
more or less movable contact with one another and with 
other parts and organs. Stimuli from the environment 
acting upon the skin receptors give rise to reflexes which 
put muscles into activity, and these activities serve in turn 
as stimuli to receptors in the deep parts ; and the impulses 
arising from these deep receptors may pass to still other 
muscles over the same effector neurons used by impulses 
coming from skin receptors. The proprio-ceptive field is 
a sort of relay and supplement and extension of the field 
of contact w T ith the environment, or extero-ceptive field. 

A simile used by Sherrington helps us to understand the 
import of this compounding of reflexes : "The receptor 
system bears, therefore, to the efferent paths the relation 
of the wide ingress of a funnel to the narrow egress. Fur- 
ther, each receptor stands in connection not with one efferent 
only but with many perhaps with all though as to some 
of these only through synapses of high resistance. The 
simile to a funnel will therefore be bettered by supposing 
that within the general systemic funnel, of which the base 
is five times wider than the egress, the conducting paths 
from each receptor may be represented as a funnel inverted 
so that its wider end is more or less coextensive with the 
whole plane of emergence of the final common paths. This 
gives some idea of the enormous formation of common 
paths from tributary paths which must take place." 10 

And Sherrington forces home the truth of the scope of 
the combinations by calling attention to the fact that under 
strychnine poisoning "a muscle can be excited from prac- 
tically any afferent nerve in the body." This is equiva- 
lent to saying, he remarks, "that each final common path is 
in connection with practically each one of all the receptors 

174 The Unity of the Organism 

of the body." 11 

Even though this statement should prove to be too strong, 
it certainly contains truth enough to show "the profusion 
in which common paths exist." 

The functional side of the total reflex system, the struc- 
tural side of which was visualized by the funnel simile stated 
above, may be regarded as pretty well presented by certain 
phenomena known as irradiation. What is meant by this 
is quite clear, in its general outlines, from the following: 

"The more intense the spinal reflex . . . the wider, as a 
general rule, the extent to which the motor discharge spreads 
around its focal area. Thus, as stimulation of the planta 
causing the flexion-reflex is increased there is added to the 
flexion of the homonymous hind limb extension of the crossed 
hind limb, then in the homonymous fore limb extension at 
elbow and retraction at shoulder, then at the crossed fore 
limb flexion at elbow, extension at the wrist, and some 
protraction at shoulder ; also turning of the head toward 
the homonymous side, and often opening of the mouth, 
also lateral deviation of the tail. According to circum- 
stance, especially according to intensity of stimulation, the 
field of end-effect of the flexion-reflex may vary from a 
minute field occupying part of a flexor muscle of the knee 
to a field including musculature in all four limbs and neck 
and head and tail." 12 

Antagonistic Reflexes in Skeletal Muscle Groups Finally 


So far what has been said about the interaction between 
reflexes has dealt only with interactions which are har- 
monious with one another in various modes of combination. 
But there are antagonistic or competitive as well as har- 
monious interactions. These must now receive attention. 
As an illustration we make use again of the scratching re- 

Neural Integration 175 

flex. "If," says Sherrington, "while stimulation of the skin 

/ j 

of the shoulder is evoking the scratch-reflex, the skin of 
the hind foot of the same side is stimulated, the scratching 
may be arrested." 13 

Then the author proceeds to show, by description and 
diagrams, how the two excitations here involved, one from 
the skin of the side of the body, the other from the hind 
foot of the same side, have the "same final common path" 
to the muscles of the hip and leg, which, however, they "use 
to different effect"; that is, the one to excite, the other to 
inhibit, contraction of the muscles concerned. 

At once there arises the important question : What is the 
meaning of such antagonisms, such seeming want of har- 
mony, as this? How deep-seated is the competition thus 
shown? Does it amount to an "ultimate truth" in the na- 
ture of the organism, thereby furnishing an argument in 
favor of the "struggle of the parts" as an explanation of 
the organism? Or is it possible that the antagonism is 
secondary to the wider needs and activities of the organism 
as a whole? 

A partial answer to this inquiry is found in the character 
of the stimuli operative in the two opposing fields. "Stimu- 
lation of the skin of the hind foot by any of the various 
stimuli that have the character of threatening the part with 
damage causes the leg to be flexed, drawing the foot up 
by steady maintained contraction of the flexors of the ankle, 
knee and hip." 

In other connections Sherrington dwells on the peculiari- 
ties of effect and importance from the standpoint of adap- 
tation, of stimuli of this class, as for example pricking, 
strong squeezing, injurious heat, and so on. 

From the descriptions we notice that the actions of the 
hind leg involved in the scratching-reflex are considerably 
different from those involved in withdrawing the hind foot 
from stimuli of harmful portent. But since the same mus- 

176 The Unity of the Organism 

cles are necessarily involved in the two sorts of action, and 
since the stimuli reaching these muscles from the two 
sources must use the same final paths to those muscles, back 
of the antagonism between the two sets of reflexes is the 
question, which, in a given case, is more important to the 
organism, the scratching action or the withdrawal-from- 
danger action? 

Since the scratch-reflex in the dog is probably connected 
primarily with flea and other insect bites, and since on the 
whole it may be assumed that these are rather annoyances 
than real dangers to life and limb as the "nocuous" excita- 
tions are by fundamental nature, it would be fair to infer 
that though either reflex might under certain circumstances 
inhibit the other, the threshold for the injury-escaping re- 
flex would be lower. I do not know that there is anything 
in the evidence which bears directly on this point, but the 
question is one that would surely arise were the whole sub- 
ject of antagonistic reflexes to be looked at from the stand- 
point of the needs and adaptations of the normally living 

The way certain other reflexes, antagonistic in a sense, 
are yet correlated in a larger sense, is more obvious than in 
the case just given. Thus the reflex appertaining to two 
limb muscles which oppose each other does not merely ac- 
tivate the muscle which contracts ; it simultaneously causes 
depression of the opposing muscle. Cooperative antagon- 
ism, as it might be called, of this general sort is widespread 
among higher animals, and applies to glandular, circulatory 
and various other mechanisms as well as to the muscular. 
Indeed, it being undoubtedly true, as Sherrington repeatedly 
points out, that the "outward behavior" of animals involves 
a great variety of movements which proceed in an orderly 
sequence, if they are normal, it seems almost necessary to 
suppose that really all normal reflexes must be cooperative 
and harmonious with reference to the organism as a whole, 

Neural Integration 177 

even though when viewed one by one or in secondary groups 
they are antagonistic. 

Such a conception of the real nature of antagonistic re- 
flexes is favored by the seemingly general fact that these 
reflexes are seldom if ever really destructive of one another, 
since they do not act upon one another simultaneously. 
Their antagonism consists in a successional opposition to 
one another. As they follow one another one acts in the 
opposite direction to the other, and the antagonism is the 
more real in that frequently they overlap to some extent. 
But as already said this overlapping probably never amounts 
to complete coincidence. Such overlappings and other 
forms of partial opposition constitute the phenomena of in- 
hibition which play a great role in the sum total of reflexes 
of the organism. This is part of the method by which 
transition is accomplished from one reflex to another, where 
the same muscles, for example, execute both. But the fact 
that the transition is accomplished normally "without con- 
fusion," to use Sherrington's phrase, shows the subordina^ 
tion of the inhibitions to the organism as a whole. 

Another important fact to which Sherrington calls at- 
tention is that inhibitions which reflexes produce upon one 
another never, so far as is known, result in injury to the 
tissues involved. Genuine opposition of reflexes, as of any 
other sorts of physical or chemical action, would, according 
to all our conceptions of natural bodies, have deleterious 
effects on the opposing bodies. As a matter of fact, in- 
hibiting reflexes not only do not injure the mechanisms in- 
volved, but actually prepare them for greater functional 
activity later on. 14 

This beneficent effect, as it might be called, of inhibition 
is perhaps illustrated by certain forms of compensatory 
reflexes. Thus, stimulation of the central end of the nerve 
to the extensor muscles of the dog's knee results in contrac- 
tion of the flexors of the hip and knee. But on removal of 

178 The Unity of the Organism 

this stimulus contraction of the extensor muscles imme- 
diately succeeds, this "rebound" being especially marked 
in the vasto-crureus muscle, the main knee extensor. 15 

The Antagonisms within the Autonomic System Finally 


It was mentioned above that "cooperative antagonism" in 
nervous action is widespread in the animal mechanism. The 
illustrations given pertained to the cerebrospinal system 
and the innervation of skeletal muscles. We must now fol- 
low this subject farther and deal with the somewhat similar 
phenomena presented by the autonomic nervous system. 
This subject was treated to some extent in the chapter on 
internal secretions, to the extent, that is, that it implicated 
the endocrinal glandular system. The presentation here 
will involve some repetition of what was said in the previous 
discussion, but the nature and importance of the subject 
will justify this. 

Cannon appears to have been the first to make clear the 
similarity between the opposing action of the subdivisions of 
the autonomic nervous system and what Sherrington calls 
the reciprocal innervation of antagonistic skeletal muscles. 
"As the above description has shown," Cannon writes, "there 
are peripheral oppositions in the viscera corresponding to 
the oppositions between flexor and extensor muscles." 

The description referred to is summed up in the state- 
ment that many of the viscera and other parts of the body 
are innervated by either the cranial or sacral, i.e., the ter- 
minal autonomies, and also by the thoracico-lumbar or mid- 
dle autonomic, this double innervation being such as to be 
statable thus: "When the mid-part meets either end-part in 
any viscus their effects are antagonistic. 9 ' The heart- 
beat is slowed by the cranial autonomic and quickened by the 
thoracico-lumbar. Contraction of the smooth muscular 

Neural Integration 179 

layer of the stomach and small intestine to produce "tone" is 
increased on the whole by the cranial autonomic, while gas- 
tric peristalsis and secretion are inhibited and the arterioles 
of these organs are contracted by the thoracico-lumbar. 11 
The pupil of the eye is contracted by the cranial autonomic 
and dilated by the thoracico-lumbar. The lower part of the 
large intestine is contracted by the sacral autonomic and is 
relaxed by the thoracico-lumbar. The discharge tube of 
the urinary bladder is relaxed by the sacral and contracted 
by the thoracico-lumbar. The blood vessels of the erectile 
tissue of the external genitals are dilated by the sacral 
autonomic and contracted by the thoracico-lumbar; and 
so on. 

Now it is especially important, as Cannon says, to notice 
the kind of service these subdivisions of the autonomic sys- 
tem perform for the organism. On considering the func- 
tions of the cranial division, one recognizes that they have 
to do with bodily conservation. "By narrowing the pupil 
of the eye they shield the retina from excessive light. By 
slowing the heart rate, they give the cardiac muscle longer 
periods for rest and invigo ration. And by providing for the 
flow of gastric juice and by supplying the muscular tone 
necessary for contraction of the alimentary canal, they 
prove fundamentally essential to the processes of proper 
digestion and absorption by which energy-yielding material 
is taken into the body and stored. To the cranial division 
of the visceral nerves, therefore, belongs the quiet service 
of building up reserves and fortifying the body against times 
of need or stress." 19 

Pasing to the sacral division, one sees that as concerns 
its distribution to the digestive and urinary viscera, its of- 
fice is that of accomplishing the discharge of refuse material 
from the body. Hence, "like the cranial division, the sacral 
is engaged in internal service to the body, in the perform- 
ance of acts leading immediately to greater comfort." 

180 Tlie Unity of the Organism 

So much in illustration of the service of the two end di- 
visions of the autonomic nervous system. 

What, exactly, we now inquire, is the nature of the service 
performed by the thoracico-lumbar division? Since this 
part is, as already seen, antagonistic in its action to both 
the cranial and sacral parts wherever it innervates an organ 
or part also innervated by these end-parts, the inference is 
readily drawn that its service to the organism as a whole 
would also be in a sense opposed to the services of the 
cranial and sacral divisions. Only a moment's reflection 
is necessary to recognize that at some points at least this 
is so. For instance, quickening of the heart beat through 
innervation by the thoracic autonomic nerves, thereby send- 
ing the blood stream more rapidly and strongly through the 
whole body, is clearly opposed, so far as these acts in them- 
selves are concerned, to the slowing of the heart and hence 
of the blood stream by the vagus nerve, i.e., by cranial au- 
tonomic innervation. Now this can mean nothing else than 
that whereas the vagal (cranial) autonomic action is in 
the interest of upbuilding and conserving the organism in 
its whole normal, wonted life, the speeded-up action through 
the thoracic autonomic is in the interest of some special, 
more or less temporary need or condition. But obvious and 
important as is this opposition between the two subdivisions 
as concerns heart innervation, of not less importance, though 
less easily observable in their full scope, are the oppositions 
and reciprocations brought about in the peripheral blood 
vessels of the whole body through the innervation of the 
muscles of the arterial walls by the thoracico-lumbar au- 
tonomic in connection with the heart action. These in- 
nervations, coupled with those of the smooth muscles of the 
gastro-intestinal canal, of the sweat and probably other 
peripheral glands, of the smooth muscles of the hairs in 
mammals, of the adrenal medulla for the secretion of 
adrenin, and of the liver for releasing stored carbohydrates 

Neural Integration 181 

into the blood, by the thoracico-lumbar autonomic, produce 
an equilibrating mechanism of well-nigh inconceivable com- 
plexity and delicacy, the workings of which are, however, 
beginning to be revealed to us, both as to details and as to 
rationale for the life of the organism as a whole. 

The whole scheme is adjusted, we may say, for these two 
necessities of the organism: (1) to serve the organism in its 
uninterrupted and uninterruptible, and therefore normal, 
growth and existence; and (2) to secure the perpetuity of 
the organism through the recurring times of special demand 
and stress which are inevitable from the external conditions 
of life under which all living beings exist. 

The supreme significance of all this from our standpoint 
is that we get a glimpse of the means by which an extensive 
and fundamental number of parts of the organism are sub- 
ordinated in their special activities to the special needs of 
the whole. The following from Cannon's chapter, Fatigue 
and Blood Pressure, gives concreteness and some particu- 
larity to this statement : "In connection with the foregoing 
considerations [of facts bearing on the value of increased 
arterial pressure in pain and strong emotion] the action of 
adrenin on the distribution of the blood in the body is highly 
interesting. By measuring alternations in the volume of 
various viscera and limbs, Oliver and Schafer proved that 
the viscera of the splanchnic area e.g., the spleen, the 
kidneys, and the intestines suffer a considerable decrease 
of volume when adrenin is administered, whereas the limbs 
into which the blood is forced from the splanchnic region 
actually increase in size. The action of the adrenin indicates 
the relative degrees of sympathetic innervation." 21 

This last sentence is, of course, what specially concerns 
us in this discussion. Continuing, and having in view his 
own researches into the beneficial effects on fatigued striated 
muscles of adrenin in the blood, Cannon says : "At times of 
pain and excitement sympathetic discharges, probably aided 

The Unity of the Organism 

by the adrenal secretion simultaneously liberated, will drive 
the blood out of the vegetative organs of the interior, which 
serve the routine needs of the body, into the skeletal muscles 
which have to meet by extra action the urgent demands of 
struggle or escape. But there are exceptions to the general 
statement that by adrenin the viscera are emptied of their 
blood. It is well known that adrenin has a vasodilator, not 
a vasoconstrictor, action on the arteries of the heart; it is 
well known also that adrenin affects the vessels of the brain 
and the lungs only slightly if at all. From this evidence 
we may infer that sympathetic impulses, though causing 
constriction of the arteries of the abdominal viscera, have 
no effective influence on those of the pulmonary and intra- 
cranial areas and actually increase the blood supply to the 
heart. Thus the absolutely and immediately essential or- 
gans those the ancients called the 'tripod of life' the 
heart, the lungs, the brain (as well as its instruments, the 
skeletal muscles) are in times of excitement abundantly 
supplied with blood taken from the organs of less importance 
in critical moments." And Cannon concludes with the very 
pertinent remark: "This shifting of the blood so that there 
is an assured adequate supply to structures essential for the 
preservation of the individual may reasonably be inter- 
preted as a fact of prime biological significance." Indeed 
so ! Even more significant, I believe, than Cannon has shown 
us as we shall see in a later chapter. But the limitations 
set at the outset of this chapter, namely that of dealing 
only with neural processes strictly, especially as these mani- 
fest themselves in reflexes, permits us to go no farther than 
to call attention to the fact that while Cannon finds "the 
most significant feature" of the reactions he has studied to 
be "that they are of the nature of reflexes they are not 
willed movements, indeed they are often distressingly be- 
yond the control of the will," 23 he yet has coupled them in 
the most positive way with the emotions, especially with 

Neural Integration 183 

those of fear and rage, and with pain. Otherwise stated, he 
has coupled them, more definitely than this has been done 
before, with the conscious psychic life of the organism. 

Concluding Remarks on the Significance of Neural Integra- 
tion for the Organismal Standpoint 

Perhaps enough illustration and general discussion have 
been presented to convince the reader not only that "the 
nervous system functions as a whole" but that this func- 
tioning is strictly subservient to the needs of the organism 
as a whole, whether the normal individual, living normally, 
or the normal individual living under special stress, be con- 
sidered. It is hoped the reader will not have failed, despite 
the brevity and inadequacy of the presentation, to perceive 
the fundamental truth that the organism's totality of activi- 
ties, executed to so large an extent through the agency of the 
neural mechanism, are in turn subordinate to the needs of 
the organism as related to its natural environment. This is 
only another way of expressing the truth, which has become 
almost trite since the idea of evolution has been an essen- 
tial part of biological philosophy, that all, or at least by 
far the major part, of the activities of organisms are 

In connection with no other organ-system than the nerv- 
ous does the truth come out so patently that the unity and 
wholeness of the particular system, the unity and wholeness 
of the organism, and the adaptiveness of the organism to its 
environment are bound together inseparably; indeed, may 
almost be called different aspects of one and the same truth 
of living beings. The very conception of adaptation, at 
least as touching neural activity, seems dependent upon the 
correlatedness, the unitedness, of the differentiated parts. 
The conception of adaptation, so far at least as reflexes are 
concerned, depends essentially on the mode of relation of 

184 The Unity of the Organism 

parts. It is non-existent except through a particular form 
of integration. 

Every specific act of every part of the nervous system 
is primarily in the interest of some other part and function 
of the organism than itself. 

This is particularly true for all adaptive acts of the 

Considerations of this sort seem not only to justify, but 
to render necessary some such view of purpose in the or- 
ganism as the following: "The infinite fertility of the or- 
ganism as a field for adapted reactions has become more 
apparent. The purpose of a reflex seems as legitimate and 
urgent an object for natural inquiry as the purpose of the 
colouring of an insect or a blossom. And the importance to 
physiology is, that the reflex reaction cannot be really in- 
telligible to the physiologist until he knows its aim." 25 No 
biologist whose mind is both open and reasonably pene- 
trating will hesitate to accept these views as correct. 

What I wish to call particular attention to is that the 
last sentence quoted calls for an additional clause: "the re- 
flex reaction cannot be really intelligible to the physiologist 
until he knows its aim," and he can know its aim only by 
considering it in the light of the organism's entire complex 
of normal activities; i.e., in accordance with the conception 
of the organism as a whole. 


1. Sherrington 6 14. Sherrington 136 

2. Sherrington 12 15. Sherrington 205 

3. Sherrington 12 16. Cannon, W. B. ('16) . 35 

4. Sherrington 38 17. Cannon, W. B. ('16) 

5. Sherrington 8 18. Cannon, W. B. ('14) 

6. Donaldson 605 19. Cannon, W. B. ('16) 

7. Sherrington 16 20. Cannon, W. B. ('16) 

8. Sherrington 8 21. Cannon, W. B. ('16) 

9. Sherrington 116 22. Cannon, W. B. ('16) 

10. Sherrington 145 23. Cannon, W. B. ('16) 

11. Sherrington 146 24. Sherrington 114 

12. Sherrington 151 25. Sherrington 236 

13. Sherrington 136 








Chapter XXI 


HAVING now carried our study of the nervous sys- 
tem far enough to enforce the conception that the 
very essence of this system, even the wholly reflex aspect of 
it, is its unifying, its integrating office for the organism, it 
remains to see what the elementalist mode of reasoning can 
do with the system in this aspect of it. 

As in the case of internal secretions, we can get at this 
subject in no way better than through the writings of 
Jacques Loeb. This way of approach will be seen to be the 
more advantageous when Loeb's long-continued and dis- 
tinguished experimental studies on some of the activities of 
several kinds of lower animals are borne in mind. 

Neglect of the Works of Sherrington and Cannon by 

Jacques Loeb 

Despite Loeb's express statement in the preface to The 
Organism as a Whole that the volume is intended to be a 
companion to The Comparative Physiology of the Brain, on 
which account a discussion of the central nervous system is 
omitted from The Organism as a Whole, the omission seems 
very unfortunate. Much important work on the physiology 
of the nervous system, especially on its integrating function, 
has been done since The Comparative Physiology of the 


186 The Unity of the Organism 

Bra'm was published,* Sherrington's really epochal book 
being especially notable among these later productions. 
Some of the most fundamental views and arguments pre- 
sented by Sherrington and others are irreconcilable with 
conceptions which are at the very center of the elementalist 
philosophy, and which find repeated expression in Loeb's 
writings on the nervous system. It is, consequently, a mat- 
ter of surprise and disappointment that a volume devoted 
expressly to the organism as a whole should leave almost 
unmentioned the very organ-system which is most distinctive 
of the organism thus viewed, especially since this leaves un- 
touched clearly formulated theoretical issues of cardinal 

Likewise the disposal of Cannon's work, upon which we 
have drawn so largely in the preceding chapter, by five 
lines in The Organism as a Whole l is truly astonishing ! 

Indeed, so striking is the defectiveness of The Organism 
as a Whole in this respect that one can not help recognizing 
that despite its author's statement in the preface as to his 
intentions, the omission amounts to an evasion even though 
not so intended. 

However, the expressed statement by Loeb of the place he 
wishes the Comparative Physiology of the Brain to hold 
in his interpretation of the organism has great usefulness 
to us : it is tantamount to an assurance that the views set 
forth in the Comparative Physiology have undergone no 
essential change because of the discoveries and arguments 
produced since the publication of that work. So our exam- 
ination of Loeb's position on this important matter must 
concern itself largely with facts and views contained in 
the Comparative Physiology of the Brain. 

The copyright of Comparative Physiology of the Brain bears the 
date 1900, while The Integrative Action of the Nervons System, by 
Sherrington, than which few more important books on the system have 
been written, was copyrighted in 1906. 

Implications of the Theories of Nerve Action 187 

The Real Importance of Loeb's Conception of the Nervous 


First of all, it is with genuine satisfaction that I recognize 
the eminent service rendered physiology and general biology 
by this author's experimental investigations on animal ac- 
tivities. I have long considered that one of physiology's 
foremost achievements is the clarification of the conception 
that nerve phenomena, the most characteristic features of 
which are response to stimuli and the conduction of the 
effects of stimulation, are not wholly unique attributes of 
nerve tissue, but are elaborations of attributes inherent in 
all protoplasm. Perhaps no single physiologist has con- 
tributed more to this clarification than has Loeb. Par- 
ticularly important has been his demonstration that nerve 
centers in the sense of ganglionic masses which determine 
reflexes are not fundamental to the coordinated and adapted 
movements of animals ; that reflexes take place in many 
animals normally and in many others experimentally, where 
no sucli centers exist, as for example in the ascidian after 
the single ganglionic mass is cut out. 

The pertinacity and technical skill with which Loeb has 
followed up these ideas are highly commendable. By focus- 
sing attention on the fact that since plants have no nervous 
system, they are, in so far as they exhibit movement induced 
by stimuli, illustrations of the principle of protoplasmic 
response in actually differentiated organic beings ; and by 
carrying the same conception over into the animal world and 
demonstrating that many activities here also are dependent 
upon direct protoplasmic response and not necessarily on 
a specially differentiated ganglionic mass, he was led to 
formulate the tropism theory. This theory, as we shall 
show presently, has a significance which Loeb himself seems 
not to have fully comprehended, his general standpoint be- 
ing unconductive to such comprehension. The theory must, 

188 The Unity of the Organism 

consequently, hold a large place in our effort to see what Loeb 
as a representative of the elementalist school in biology is 
able to do with the integrative action of the nervous system. 
Let it be understood that with the general controversy 
which has gone on so long and warmly concerning the value 
of the tropism theory for explaining animal behavior, we are 
little if at all concerned in this discussion. The point of 
cardinal importance for us, and the thing about the theory 
which, as mentioned above, Loeb and his followers seem not 
to have perceived clearly, is the fact that the theory is 
really dependent for so much of validity as it has on the 
conception of the organism as a whole. In other words, the 
theory is in essence an organismal and not an elementalistic 
theory. Hence it results that since Loeb is the author and 
chief proponent of the tropism theory, but is at the same 
time an extreme elementalist, his general position as an in- 
terpreter of animal activities and the nervous system con- 
tains fundamental contradictions. Indicating these con- 
tradictions and exploring, though incompletely, the course 
along which the truth lies, will form a natural conclusion to 
our study of the integrative office of the nervous system. 

The Organismal Character of Tropisms Partly Recognized 

By Loeb 

I first call attention to the fact that Loeb himself has seen, 
though "as in a glass darkly," the organismal character of 
the tropism theory. Speaking of the lack of a sharp boun- 
dary between reflexes and instincts, he says some authors 
are wont to speak of reflexes when the responses to stimuli 
pertain to single parts or organs, but they "speak of in- 
stincts where the reactions of the animal as a whole are 
involved (as is the case in tropisms)." 2 

And the same idea comes out still more definitely in a 
proposal to discard the term reflex as applied to such a re- 

Implications of the Theories of Nerve Action 189 

sponse as the clasping act by the arms of the decerebrated 
male frog, and substituting the term tropism. "It is better,'' 
he says, "to call them tropisms since the organism as a whole 
is involved." 3 

But Loeb's most illuminating recognition of the organis- 
mal character of tropisms is found not in any positive 
acknowledgements but in his statements of what a tropism 
consists of essentially. This he has told us many times and 
in varied phraseology during the last twenty-five years. The 
following from The Organism as a Whole will serve well as 
the basis of our consideration: "Animals possess photo- 
sensitive elements on the surface of their bodies, in the eyes, 
or occasionally also in epithelial cells of their skin. These 
photosensitive elements are arranged symmetrically in the 
body and through nerves are connected with symmetrical 
groups of muscles. The light causes chemical changes in 
the eyes (or the photosensitive elements of the skin). The 
mass of photochemical reaction products formed in the 
retina (or its homologues) influences the central nervous sys- 
tem and through this the tension or energy production of 
the muscles. If the rate of photochemical reaction is equal 
in both eyes this effect on the symmetrical muscles is equal, 
and the muscles of both sides of the body work with equal 
energy ; as a consequence the animal will not be deviated 
from the direction in which it was moving. This happens 
when the axis or plane of symmetry of the animal goes 
through the source of light, provided only one source of 
light be present." 4 

Reduced to its lowest terms, the tropistic mechanism de- 
scribed here and in many other places consists of: first, 
portions of the body surface having specific irritability, as 
receptor organs, these being usually paired and usually 
symmetrical, though often unsymmetrical ; second, muscles 
as effectors, almost always paired and usually symmetrical; 
third, nerves, variously constituted, as conducting paths 

190 The Unity of the Organism 

connecting more or less directly the sensory areas with the 
muscles. In a word, tropistic movements are determined by 
a definite number of definite kinds of parts or organs, defi- 
nitely arranged with reference to one another and to the 
whole ; that is, upon an organized body called a living being, 
or briefly an organism. A morphological entity is funda- 
mental to the tropism conception. Tropisms are explained, 
partly, by these organizations. Loeb does not avoid, in fact 
at times appears not to try to avoid, recognizing this : "The 
irritable structures at the surface of the body and the ar- 
rangement of the muscles determine the character of the 
reflex act." "The explanation of them [tropisms] depends 
first upon the specific irritability of certain elements of the 
body-surface, and, second, upon the relations of symmetry 
of the body." 5 

The Organismal Character of the Segmented Theory of the 

Nervous System 

Another province in which Loeb has done distinguished 
work and into which the organism as such constantly ob- 
trudes itself and will not accept relegation to a secondary 
place, is that of the coordinated action of the central nerv- 
ous system in those animals in which that system consists 
of an axial series of ganglia. Here again, as in the tropism 
theory, the basal perception underlying the segmental theory 
of nerve physiology is the truth that nerve "centers" as the 
"seats" of particular activities and functions in the old sense 
(in the sense, that is, of being the real source of these 
activities) do not exist. According to the segmental theory 
the real unit of activity of an animal made up of metameres 
or segments is the segment itself, the nerve ganglion of the 
segment being only one element in the complex. The idea 
is set forth with fullness and perspicacity in the chapters 
of the Physiology of the Brain dealing with worms and 

Implications of the Theories of Nerve Action 191 

arthropods, and especially in the one on the segmental the- 
ory in vertebrates. 

The starting-point chosen by Loeb for the treatment is 
the earthworm. It is generally known that this animal is 
jointed, or segmented, that every segment has a pair of nerve 
ganglia situated underneath the intestine on the floor of 
the body wall, and that the pairs of ganglia are all connected 
with one another fore-and-aft, so that the whole constitutes 
a chain of ganglia extending the entire length of the body. 
In addition to this series of ventral ganglia, there is a pair 
of large ganglia above the gullet at the extreme front end 
of the creature, these being connected with the ventral nerve 
chain by commissures. These dorsal nerve masses are usu- 
ally called the supra-esophageal ganglia, but are often 
spoken of as the brain. 

On the basis of the "center" theory of the nervous sys- 
tem, it was very natural to raise the query: Is the earth- 
worm's brain the "seat", the primal source of the coordi- 
nated activities, the crawling and burrowing which make 
up so large a part of the life activities of these crea- 
tures? Loeb's own formulation of the question is good. 
"Does coordinated progressive movement in which all the 
segments of the body participate, depend upon the brain?" 

The first attempt to answer this question experimentally 
seems to have been made by Friedlander. Having made a 
small opening in the side of the body of a living worm and 
cut out a piece of the nerve cord so that a complete hiatus 
in the cord was produced, the author fully expected, he tells 
us, that after the healing of the wound in the body wall, 
which took place very quickly and completely, the parts of 
the body in front of and behind the section of the cord would 
either behave like two separated individuals, or that the 
front part would draw the rear part passively about. But 
neither result followed. "Animals from the middle of which 
a piece of the central nervous system is wholly wanting, 

The Unity of the Organism 

crawl exactly like normal animals except as to a small dif- 
ference to be noticed later." The wave of muscle con- 
traction and other movements, starting at the anterior end, 
passed over the point at which the ganglionic chain was in- 
terrupted and on to the hind end of the body just as though 
there were no such interruption. "The movements were 
coordinated exactly in the same way as in the normal ani- 

These experiments, together with others that have since 
been performed by other investigators, overthrow, as Loeb 
says, "the idea that coordination in these animals is deter- 
mined by a special centre of coordination which is located 
in the brain." 7 

But if the brain is not the coordinating* center and does 


not contain it, what and where is that center? The answer is 
there is no such center. Coordination is not a central matter 
at all. Rather it is a matter of the working together of 
many parts, each duly balanced off with many others, and 
properly subordinated to the whole animal. 

The details of the truth thus stated in general terms can 
be illustrated more advantageously by referring to some of 
Loeb's own observations on a group of annelids considerably 
higher in the scale of life than the earthworm. I refer to 
the pile worms familiar to many persons accumstomed to 
marine shore fishing. This group of worms, known collec- 
tively as nereid annelids, are far more "heady" than earth- 
worms in that the useful body member named head is more 
definitely set off from the rest of the body than it is in the 
earthworm, and is outfitted with eye-spots, touch appen- 
dages, and so on, wholly lacking in the earthworm. Nereis 
is much more highly "cephalized" than is the earthworm, so 
brain centralization would be expected to present features 
here not occurring in the more lowly worm. When nereids 
are deprived of the supra-esophageal ganglia their behavior 
is such as to suggest at first sight support for the concep- 

Implications of the Theories of Nerve Action 193 

tion that the brain is a fundamentally originative and coor- 
dinative member. For example, an individual thus depleted 
will crawl about unceasingly on the sand of the aquarium 
bottom, without, however, showing any signs of burrowing 
into the sand, though this operation is very characteristic 
of the normal animal. This looks as if the brain were indeed 
the seat of the burrowing instinct. But not so. The bur- 
rowing activity can be induced not only in animals deprived 
of the brain, but even in parts of animals from which the 
heads have been cut away altogether. Loeb placed a brain- 
less piece of a Nereis on the sand, and as usual it remained 
quiet. "I then gradually covered the forward end with 
sand. The rest of the animal immediately began to make 
the typical movements which the animal makes in forcing its 
way into the sand. At the same time the glands began to 
secrete the sticky substance which cements the particles 
of sand together, forming the wall of the burrow-hole." 
"But, why," Loeb asks, "does the Nereis not burrow when 
deprived of its brain? For the simple reason that it makes 
use of the organs of the mouth in burrowing, and these are 
amputated with the head. Hence it is the loss of the pe- 
ripheral head-organs which keeps the decapitated Nereis from 
burrowing, and not the loss of the brain." 

Since the coordinated activities involved in burrowing can 
be accomplished through the acting together of various su- 
perficial and deeper body parts sense organs, muscles, and 
so on without the intermediation of the brain, these pe- 
ripheral members are the real "seat" of such activities. The 
author then goes on to inquire what part the brain does play 
in the creature's normal life. "If we compare," he says, 
"the conduct of a Nereis whose brain has been amputated 
with that of a normal worm, the difference seems to be of the 
same nature as that between an insane and a rational human 
being. . . . The peculiar irritability by means of which the 
Nereis draws its head back and moves backward out of the 

194 The Unity of the Organism 

tube depends upon organs which are located in the forward 
end of the body and whose sensory nerves go to the supra- 
esophageal ganglion." 

One office of the Nereis brain is, therefore, to make a 
great complex of activities normal, or "of the same nature" 
as sane activities in "rational human beings." Not the ac- 
tivities themselves but their normality are dependent on the 
brain. This distinction is undoubtedly general, but it is 
definite so far as it goes. Beyond question to the student 
who should compare the movements of the normal and the 
brainless Nereis with sufficient care, the distinction between 
the two would be as positive, as indubitable, and as definable 
as that between almost any two complexes of organic phe- 
nomena. The unsatisfactoriness of the distinction is not its 
vagueness but its generalness. It is sufficiently definite 
though not sufficiently analytic. 

Just what the brain does to normalize the activities ; in 
other words, how the change of character of the worm's 
behavior its "restlessness" and general insane-like conduct 
results from the absence of the brain, is not known. Loeb 
makes a suggestion toward an explanation, but recognizes 
what he offers as nothing more than a suggestion ; so to go 
into it is not w r orth our while. 

But there is an obvious usefulness of the ganglia and 
whole neural chain in such animals aside from the general 
one of normalizing behavior. It serves as a common con- 
ducting and relay system between the peripheral sense or- 
gans and locomotor muscles. Loeb's statement of the matter 
is concise and may be adequate : "The central nervous sys- 
tem does not control response to stimulation ; it merely 
serves as a conductor from the point of stimulation to the 
muscle through which weaker stimuli may pass, and pass 
more rapidly than would be possible if the muscle were stim- 
ulated directly. 

"In the Annelids each ganglion is a relay station for 

Implications of the Theories of Nerve Action 195 

the sensory and motor nerves of the corresponding segment. 
If the head exercises a stronger influence upon the behavior 
of the animal than any other segment, as in Nereis, for in- 
stance, I believe it is due to the fact that in the oral end 
more kinds of irritability are present and more peripheral 
organs are differentiated (sense organs, mouth, etc.) than in 
the other segments." 10 

Loeb has done good service in bringing together the evi- 
dence for, and emphasizing the truth that, the segmental 
make-up of the vertebrate head, so long and so ardently 
studied by anatomists and embryologists, is as important a 
matter for neural physiology as it is for morphology, and 
that as a matter of fact the vertebrate spinal cord works 
more or less on the segmental principle, and in this respect 
falls into line with the central nerve axis of annelids and 

The first, or one of the first, investigators to recognize 
this truth was Schrader, in his studies on the activities of 
frogs whose brains had been wholly or partly removed. The 
frequently quoted and later fully confirmed statement by 
this investigator is of prime significance for the general 
truth of what we might call fore-and-aft neural integration 
in vertebrates. "The series of experiments we have given 
teaches us that the central nervous system of the frog can be 
divided into a series of sections, each of which is capable of 
performing an independent function. It brings the central 
nervous system of the frog into closer relation with the cen- 
tral nervous system of the lower forms, which consist of a 
series of distinct ganglia that are connected by commissures. 
It speaks against the absolute monarchy of a single central 
apparatus and against the existence of different kinds of 
centers, and invites us to seek for the centralization in the 
many-sided coupling of relatively independent stations." 

As statements much at variance from this concerning the 
"central" function and power of different parts of the 

196 The Unity of the Organism 

frog's brain are not uncommon, and as the facts are spe- 
cially important for u.s, it is desirable to consider the evi- 
dence a little more in detail. To this end the following from 
Holmes may be taken as an authoritative summing up of 
results so far reached in this field. "A frog; with the brain 


removed as far back as the medulla is still capable of per- 
forming regular leaps and swimming movements of the 
limbs. When thrown on its back it rights itself, and still 
performs compensatory motions when tilted or rotated. 
Breathing is normal, and the animal swallows pieces of food 
that are placed in its mouth." 12 

If the removed portion of the brain is made yet larger, 
by including the anterior part of the medulla, the animal is 
still capable of performing a long series of regular activi- 
ties. "Locomotion is effected mainly by creeping, but the 
frog is nevertheless capable of springing in the ordinary 
manner. In the water it swims by alternate movements of 
the limbs. . . . The breathing and swallowing reflexes are 
still normal, but the croaking reflex is no longer performed. 
The reflex of snapping at food is not destroyed. ... If a 
piece of meat is rubbed against the frog's nose, the animal 
snaps at it and uses the fore legs to stuff it into the mouth. 
The same reaction may be brought about by using the finger, 
but after the finger is seized and it is found that the object 
is too large to be stuffed into the mouth, the frog begins to 
reject the morsel, and uses the fore legs to push it away. 
Truly a remarkable combination of reflexes!" 12 

And further : "There is no center for coordinated locomo- 
tion in the medulla. Disturbances of locomotion begin with 
the fore limbs. If the medulla is cut across at the tip of 
the calamus scriptorius, the animal sinks on its breast, and 
the fore limbs are for a considerable time helpless, although 
the hind limbs are capable of performing vigorous coor- 
dinated movements. The reason for this is that the injury 
lies so near the region from which the nerves of the fore 

Implications of the Theories of Nerve Action 197 

limbs arise that the movements of these members are very nat- 
urally affected." 13 It is the interdependence fore-and-aft 
of reflex activities like those here indicated, that is, of the 
fore limbs acting with the mouth, and the hind limbs with 
the fore limbs, that suggested to Loeb the idea of "chain 
reflexes," a conception of special interest from our stand- 

That the segmental mode of functioning of the vertebrate 
nerve axis is not limited to the lower portion of the great 
vertebrate phylum is, as Loeb points out, shown by such 
investigations as those of Goltz, and of Goltz and Ewald on 
the dog. These are given so fully in the larger text-books 
of physiology and are thereby so readily accessible to all 
specially interested in the subject, that a very brief presen- 
tation will suffice. 

Two main groups of reflex centers in the nerve cord are 
made out, the cervico-thoracic and the lumbo-sacral. The 
first group contains the center for movements of the an- 
terior limbs, for respiratory movements, for acceleration 
of heart action, for dilation of the pupil of the eye, and for 
several other activities at the anterior end of the body. The 
lumbo-sacral group contains the center of movements for 
the posterior limbs, for control of the anal sphincter, the 
activities of the generative organs, and several other parts. 

The existence of such centers in the dog's cord w r as proved 
by section of the cord at different levels under the most 
careful operative conditions. Had the investigation stopped 
with experiments of this sort the facts brought out might 
have been interpreted as confirmatory of the "seat" notion 
of a nerve center. When, however, it was discovered that a 
dog from which most of the cord is removed may live for 
many months in a good state of health, all the vegetative 
functions being carried on almost typically, it became clear 
that the various centers of the cord are not the seats of life 
activities in any such fundamental sense as was earlier 

198 The Unity of the Organism 


For one thing the results seemed to show that the sym- 
pathetic nervous system shares with the central system 
more profoundly than had been known before, in determin- 
ing life processes. 

The conclusions which flow with apparent certainty from 
the observations are well summed up in the following: "Un- 
doubtedly all such activities [of visceral life] may subsist 
and function in a comparatively normal fashion after re- 
moval of all spinal influence. The office of the spinal sys- 
tem in regard to the visceral life seems to consist in en- 
dowing these functions with greater energy, and in con- 
ferring greater stability and more solid equilibrium on the 
general constitution of the animal." 14 

The reader will hardly fail to recall, on reading this state- 
ment about the function of the mammalian spinal cord, 
Loeb's statement, quoted some pages back, about the nor- 
malizing function of the Nereis brain. "Stabilizing" the 
dog's function means much the same as "normalizing" the 
annelid's functions. Nor should the reader neglect to notice 
that he might substitute the word "integratedness" for 
"equilibrium" in the quotation without change in the es- 
sential meaning of the sentence. 

Critique of the Elementalist Attempt to Interpret Tropistic 
and Segment al Theories of the Function of the 

Nervous System 

We now return to the central point of the present sub- 
topic, namely that of what a genuine bio-elementalist is able 
to do when confronted by the facts in possession of modern 
physiology bearing on the tropistic and segmental theories 
of the nervous system. 

First we are compelled by the evidence to recognize the 
general soundness of the doctrines. Second, we recognize 

Implications of the Theories of Nerve Action 199 

that probably no biologist has dealt with the conception 
more comprehensively and illuminatingly than Loeb. Third, 
we find that his espousal and able treatment of the theories 
has led him into positions so thoroughly organismal in both 
essence and expression as to be quite irreconcilable with his 
own more general elementalistic philosophy. 

It remains to point out more specifically than we have 
wherein the organismal implications of his teachings as ex- 
emplified by his writings constitute a refutation of the 
elementalistic implications of his teachings as exemplified 
by his strivings after an "ultimate explanation" of organic 
phenomena in the terms of physics and chemistry. 

The essence of the irreconcilability of the two positions 
may be put into a general form thus : in innumerable state- 
ments and definitions found in his discussions of these the- 
ories, Loeb is compelled to introduce the organism either as 
a whole or in considerable portions, as a causal explanation 
of particular phenomena with which he deals ; and the com- 
pulsion to such introduction makes it impossible for him or 
any one else to dispense with the causes thus introduced by 
resolving them into ultimate elements of any sort, whether 
organic or inorganic. 

Illustration and justification of this general statement 
must be given. The following typical sentence may intro- 
duce the discussion: "The irritable structures at the sur- 
face of the body, and the arrangement of the* muscles, de- 
termine the character of the reflex act." 15 

Notice what it is that "determines" the character of the 
act. "Structures" and "muscles" do it, these being "ar- 
ranged" so-and-so, the irritable structures definitely on the 
surface of the body and the muscles within the body. The 
point needing special attention is that not chemical com- 
pounds or even living substances but structures organs 
and these arranged; that is, entities which neither exist nor 
can exist except through the agency of an organism, enter 

200 The Unity of the Organism 

into the definition. 

That the structures referred to and all others of the or- 
ganism are composed of chemical substances and nothing 
else, is beyond question. Why, then, does the author not 
bring into the definition this truth about the ultimate make- 
up of the structures? The central aim of his researches 
on animal activities being what they are "a physico- 
chemical analysis of behavior," is it not surprising that he 
should be satisfied with even a working description or defini- 
tion of a reflex act which takes no notice of the results of 
such analysis? If the understanding of complicated life 
phenomena consists in resolving them into their simple ele- 
mentary components (see the first sentence of the Physi- 
ology of the Brain), then a definition of a tropism that 
would contribute largely to an understanding of it should 
not be content with such proximate constituents as "irri- 
table structures," "muscles," and so on, but should go right 
back to the ultimate physico-chemical elements. 

The typical elementalist answer to these restrictive criti- 
cisms is well known. It is, substantially, that the understand- 
ing and hence the definition of tropism, or for that matter 
of any other life phenomenon, is final or ultimate only when 
expressed in physico-chemical terms. The criticism sug- 
gested concerning the mere proximateness and hence inade- 
quacy of a definition that uses such terms as "structures," 
"muscles" aiad "body surface" is allowed to have a large 
measure of validity, the usage being justified mainly, it is 
claimed, on practical grounds. The morphological concep- 
tions involved, it is held, are so strongly intrenched in bio- 
logical terminology, indeed are so necessary in a historic 
and subsidiary sense, that it is very inconvenient, if indeed it 
is not impossible, to dispense with them. In a word, the 
contention is that while morphology and general physiology 
are necessary, their necessity is secondary or subsidiary to 
physics and chemistry. But what should be seen in this 

Implications of the Theories of Nerve Action 201 

with the greatest possible clarity is the error, the fallacy, not 
of objective fact but of reasoning. Stated in general terms, 
the fallacy consists in the tacit assumption that the indis- 
pensability and adequacy of physico-chemical elements to 
explain organic behavior are of a sort that exclude the in- 
dispensability and adequacy of "structures," as sense or- 
gans and muscles, from the explanation of the same phe- 

To go into the epistemological and logical necessities 
involved in the situation with which we are here confronted 
is entirely beyond the scope of this volume. However, in 
the interest not so much of Truth in the abstract as of 
healthy, wholesome, useful science, biologists will have to 
cease employing such deprecatory epithets as "anthropo- 
morphism," "metaphysics," "rhetoric" and the like to gain 
for themselves a sense of security in the use of language and 
reasoning which can not endure for an hour the searchlight 
of really careful thinking and expression. 

To focus the general statement of the fallacy on the par- 
ticular matter in hand, one must see that both factually 
and epistemologically the organs and other morphological 
and general functional elements, or factors found by the 
analysis of the organism, are "fundamental" or "ultimate" 
for the phenomena to be explained by exactly the same cri- 
teria that the physico-chemical elements are fundamental 
or ultimate. 

A reflex act or a trophism can no more be intelligibly 
expressed or understood or conceived as an objective fact 
without sense organs, muscles, etc., than without physico- 
chemical substances. If one questions the truth of this af- 
firmation let him test the matter by trying to express a re- 
flex act in the terms of the physico-chemical elements known 
from analysis to be "behind" such an act. To begin with, he 
finds it necessary to fix upon some particular reflex act, 
scratch-reflex, perhaps. Such particular act must be taken 

The Unity of the Organism 

as the starting point, otherwise the experimenter will be com- 
mitted to the Idea of the act rather than the act itself. 
And I take it for granted that no truly present-day biologist 
is willing, even though he may not be able to justify fully his 
unwillingness, to commit himself to that ancient position. 
Very well then, the scratch-reflex is going to be expressed in 
terms of the oxygen, nitrogen, carbon, phosphorus, and so 
on, which we are sure are "behind" it. Having recognized 
the necessity of starting with some particular reflex act, and 
having settled upon the scratch-reflex, true to his recogni- 
tion of the necessity for particularity he sees he must go 
still further in the same direction. If his analysis is to be 
exhaustive and his expression adequate, and final, it will 
have to particularize still further and concern itself with 
the reflex of some particular animal species, very likely even 
with some single individual or small group of individuals. 
A vast mass of evidence makes it almost certain that a 
dog's scratch-reflex is different from a cat's, and both are 
different from an ox's, a frog's, and so on. Suppose, then, 
the dog's reflex settled upon, and defined so fully that it 
is distinguished from every other reflex whatever. The stu- 
dent is now ready for his main undertaking, that of ex- 
pressing the dog's scratch-reflex in terms of the physico- 
chemical elements to which the reflex is reducible. 

The next step is to examine the chemical elements con- 
cerned, as these are treated in inorganic chemistry, for the 
purpose of seeing what their "terms" are; that is, what 
their attributes or properties are, the special purpose of this 
examination being to ascertain what terms, i.e., what at- 
tributes, answer to, or correspond to, and so can be used to 
express, the dog's scratch-reflex. The outcome of this 
examination is unequivocal. It finds no terms, no attri- 
butes whatever, which by themselves suggest even remotely 
the reflex under consideration. 

But since the examiner has satisfied himself that no other 

Implications of the Theories of Nerve Action 203 

simple elements, material or immaterial, are "behind" the 
reflex, and since the reflex is an indubitable reality, there is 
no escape from the conclusion that something other than the 
original inorganic simples must have intervened between 
these simples and the reflex. And what is that something? 
Does common experience and common sense hesitate with its 
answer? If it does its hesitancy is probably from surprise 
that so obvious a matter should be made the subject of 
serious questioning. "The dog is what has intervened be- 
tween the chemical simples and the reflex." That is what 
common experience must answer and will unhesitatingly 
answer once it recovers from its surprise at being ques- 
tioned on a subject so open and daylight clear. 

But then science comes forward with its criticism of this 
common-sense answer. There is no gainsaying, it admits, 
the truth of the naive answer thus given. But this answer, 
science says, is a mere truism. It leaves the case just where 
it was before science began its analysis, so is worthless for 
scientific purposes, however useful it may be for ordinary 

This rejoinder by science must be looked into carefully; 
otherwise its weakness will be missed. It must be exacted 
of science that she show more explicitly than she has what 
she means by explaining the dog's scratch-reflex by refer- 
ring it to the physico-chemical elements at the basis of the 
act. Let us, we must insist, hear you express a dog's 
scratch-reflex in the terms of oxygen, carbon, et cetera. 

That such expression is possible is freely granted. But 
how can it be done? That is a key question. It can be 
done in one and only one way, namely by adding to the 
attributes, that is, to the "terms" which inorganic chemistry 
recognizes in the chemical elements concerned, just those 
attributes and terms which the dog's scratch-reflex requires 
in order that the elements may explain the reflex. We can 
say that besides the specific gravity, combining weight, and 

The Unity of the Organism 

other well-known attributes of the elements, they possess 
dog's-scratch-reflex attributes. But, the name thus sug- 
gested for the newly discovered attributes being cumber- 
some, we may devise for them some term more simple and 
convenient for example, do-sca-re-x, doscarex. 

In virtue, then, of the doscarecious powers of oxygen, 
carbon, and so forth (the fact that these powers can only 
be assigned in the lump and not distributively to the severa] 
elements should not be lost sight of though it may be neg- 
lected for the present argument), a complete physico-chem- 
ical explanation of the phenomenon under consideration is 

Does this discussion advance the interpretation, the under- 
standing of biological phenomena beyond the familiar sar- 
casm about explaining drug-induced sleep as due to a dor- 
mitive principle of the drug? Yes, I maintain, it does, 
because in place of a "principle" attached to no particular 
drug but to any sleep-producing substance, the doscarecious 
powers recognized by us are definitely assigned to the chem- 
ical elements known to be the sole constituents of organ- 
isms. The new powers take their place perfectly, definitely 
and positively among the other attributes of the elements, 
the assignment being based on the solid ground of analyses, 
laboratory and other, made through years of scientific re- 
search. But an exceedingly important point which comes 
in sight here is that though these doscarecious powers are 
proved to be real ones, they are latent and wholly unknown 
to inorganic chemistry for the reason that thev never mani- 

o / * 

fest themselves under any other combination of things than 
just that which in its totality Zoology has named dog. A dog, 
and a dog only, is able to cause oxygen, carbon and the 
other elements to reveal these particular scratch-reflex 
powers. The dog comes in as a sine qua non to the pro- 
duction of, and hence to the causal explanation of, the 
particular group of activities under consideration. We 

Implications of the Theories of Nerve Action 205 

have no evidence that the chemical elements operating them- 
selves can actualize their own latent doscarecious powers. 

The reader will hardly fail to connect what we are say- 
ing with the familiar phenomena of the assimilation by 
organisms of nutrient substances. All our argument really 
does to the usual conception of this phenomenon is to focus 
attention upon the fundamental importance of the individual 
organism as a factor, as a cause, of the chemical transfor- 
mations wrought in the nutrient substances. The insuf- 
ficiency of statement of the assimilative, or anabolic, or 
synthetic aspect of the metabolic cycle in the organism, 
lies in its failure to bring out clearly enough the indubitable 
fact that the final results are innumerable activities and sub- 
stances which pertain solely to the living, normal individual 
w r hich are strictly personal and private, as one may say. 

The current mode of expression according to which the 
assimilative syntheses take place in the organism is quite 
misleading in that it permits or even encourages a concep- 
tion that the syntheses have a measure of detachment from, 
and independence of, the life of the organism as a working 
unit, which as a matter of fact they do not have. Again, the 
usual statement that the syntheses result in organic sub- 
stances of more complex, higher make-up is inadequate in 
that it diverts attention from the fact that these new sub- 
stances belong to, are fundamentally part and parcel of the 
organism. They are not "any old organic substances" but 
are exactly those substances necessary to maintain the nor- 
mal life of the particular individual organism. Hence de- 
spite the indubitable fact that the final results are reached 
by way of innumerable purely physical and chemical opera- 
ations, the organism itself, acting as an integer no matter 
how complex, is always to the fore as a controlling, domi- 
nating factor. 

206 The Unity of the Organism 

The Bearing of This Critique on "Analysis" in Biological 


The considerations thus briefly set forth lead to certain 
still more general ideas of the utmost importance. The 
natural entities to which we apply the descriptive terms 
living and organic are specially distinguished by the chem- 
ical and physical syntheses which they accomplish by virtue 
of their inherent constitution. So as concerns the most 
characteristic of these syntheses, especially the chemical 
ones, they are to a very considerable and fundamental ex- 
tent definitive of the organic individual, species, genus, and 
so on, of taxonomic biology. This is equivalent to saying 
that the synthetic operations regarded each by itself ter- 
minate in results which are in large measure unique and so 
unforeseeable from anything we know about the original 
elements as such; that is, before they have actually been 
subjected to the particular synthetic transformations under 

And this again is equivalent to saying that synthesis 
transformatory synthetic processes and products is more 
distinctive of living beings than are analytic products and 

Finally, it follows from these facts about the synthetic 
nature of organisms, and from the established principles of 
thought, that analysis alone is incapable of interpreting, of 
understanding organic beings. No natural object which in 
its nature is more distinctively synthetic than analytic can 
be understood by knowledge-processes which are more ana- 
lytic than synthetic. 

This conclusion goes to the very heart of the elementalistic 
position, and, as stated in the discussion on internal secre- 
tions, is really as much an epistemological as a biological 

Reverting again to Loeb's writings, the conclusion to 

Implications of the Theories of Nerve Action 07 

which we are forced joins issue with the very opening sen- 
tence of the Physiology of the Brain, as we have already in- 
dicated. "The understanding of complicated phenomena," 
particularly as presented by organisms, can not be accom- 
plished through "an analysis by which they are resolved into 
their simple elementary components." Unquestionably an- 
alysis is essential to, but it is not adequate for, full under- 
standing of the phenomena. 

The very nature of organic synthesis and of knowledge- 
getting precludes the possibility of attaining the hind and 
degree of understanding of organisms which element alls t 
biology claims to have attained and promises to attain. 

"A complete explanation of life in terms of physics and 
chemistry" is impossible for the sufficient reason that physics 
and chemistry as such do not contain any of the really dis- 
tinctive terms of life. Those terms can only be brought 
into physics and chemistry after and not before the phe- 
nomena of life have been searchingly scrutinized ; that is, an- 
alyzed and found to involve physics and chemistry. The 
terms of life are in the original data on the phenomena of 
life, and no sort of analysis can possibly make this other- 

Our position, it should be noticed, touches the hackneyed 
controversy over vitalism and materialism only in so far as 
the course of reasoning we have pursued involves the recog- 
nition that each organism (the dog, for instance) is a 
natural object possessed of certain causal powers, by ex- 
actly the same logical and epistemological criteria that any 
simple chemical element or chemical compound is a natural 

Stated in a brief and common-sense way, our contention 
is that the attributes which make a dog a living body are no 
less natural than are the attributes which make carbon a 
chemical body. The tiresome and meager-fruited contro- 
versy between Materialists and Vitalists may be character- 

208 The Unity of the Organism 

ized as due to the fact that neither party has taken the 
trouble to establish clearly, even in their own minds, the 
meaning of the word natural. As a consequence of this 
slipshodness the two groups agree tacitly, in treating the 
inorganic and the organic worlds as though natural does 
not mean the same thing in the two realms. The implica- 
tion is that if the inorganic world, for instance, be held to 
be natural by both parties, for the Vitalists the living realm 
is largely sM>7'-natural, while for the Materialists the same 
realm is largely i?zfr-natural. 

Theories of Animal Behamor in Relation to the Science of 


This somewhat protracted though wofully insufficient 
treatment of neural integration may close with a brief 
section on some of the still larger biological and methodo- 
logical implications of the conclusions reached. Special at- 
tention is called to the fact that the culminating part of 
our argument has involved data and conceptions which are 
as unequivocally zoological, morphological, and physiologi- 
cal, as any of the data and conceptions are unequivocally 
physical and chemical. Physical chemistry, or any other 
aspect of inorganic chemistry, is utterly powerless, so far 
as we can see, to discover such facts, as for example, that 
oxygen, carbon, nitrogen, etc., possess latent "doscarecious" 
powers. This final section is in the direction, consequently, 
of establishing the parity, to claim the least, of zoology? 
botany, morphology, and general physiology, with chemistry 
and physics, in the great complex group of biological 

We may first allude to a favorite mode of expression of 
materialistic element alls ts. Whenever fuller analysis lias 
proved some group of animal phenomena not hitherto con- 
nected directly with physico-chemical substances and forces, 

Implications of the Theories of Nerve Action 209 

to be in reality dependent on such agencies, this school is 
wont to remark in substance that investigation has finally 
"transferred" the phenomena from the provinces of zoologv, 
morphology, general physiology and the other sciences of 
animal life, to physics and chemistry. Our argument puts 
beyond question the logical inadequacy of such a statement. 
Analysis does not by any means transfer the phenomena 
from zoology, etc., into physics and chemistry. Neither 
analysis nor any other agency can any more take the study 
of animal phenomena away from zoology and put it into 
physics and chemistry than it can take bread-making away 
from the baker's art and put it into physics and chemistry. 
The chemist may undoubtedly take to bread-making and find 
that his new employment has much in common with his old; 
but in so far as he really succeeds at the new, he is more a 
baker than a chemist. He has not transferred bread-making 
to chemistry, but if anything has done just the reverse. 
What analysis actually docs in these cases is to extend the 
bounds of physico-chemical forces and laws into zoology* 
morphology, etc., and to prove that if zoological, morpholog- 
ical and physiological undertakings are to move into ever 
greater fullness, aid from physics and chemistry is indis- 

Thus critical examination of the reasoning of elemen- 
talist biology reveals the logical fallacy in any sort of state- 
ment which involves the assumption that the older sciences 
of organic beings, like taxonomic botany and zoology, geo- 
graphical distribution, morphology, general physiology and 
so on, are not and never can be relegated to places of minor 
or secondary importance in biology. But it is the practical 
harmfulness of such assumptions rather than the logical 
fallacies underlying them which chiefly concern us in this 
volume, and no part of our whole subject is more vitally 
affected by such harmfulness than this of the behavior, even 
the purely tropistic behavior, of animals. The whole round 

210 The Unity of the Organism 

of animal biology attitude toward research problems and 
undertakings, valuations and interest in different fields of 
knowledge, educational aims and method all are affected. 

In the light, for instance, of such a complex of animal 
behavior as that presented by the northern fur seal, its 
mating, breeding, migrating and other habits, the conten- 
tion that the myriads of complicated phenomena which go 
to make up animal life can be understood by converting 
zoology into a laboratory and experimental science, to the 
end of analyzing the phenomena into their "simple elemen- 
tary components," is so ludicrous as hardly to need argu- 
mentative refutation. Indeed, it seems as though persons 
obsessed by a theory to the extent of being impervious to 
the ludicrousness of the contention, are likely to be also 
impervious to the true reasoning involved in it. It may, I 
think, be assumed that so much of zoology as has formed 
this remarkable conception of itself will before long drop 
into the background by scientific gravitation despite its 
present great vogue. 

One of the leading motives, consequently, of this con- 
structive part of my enterprise is to establish the essen- 
tiality of general zoology and its time-sanctioned depart- 
ments on so solid a basis of philosophic reasoning that the 
necessary methodology of the regenerated science of the 
future will be clearly seen in broad outline. 

If the considerations inadequately presented in these last 
pages and in other parts of this volume once get secure 
lodgment in biological thought it will become manifest that 
the "behavior" of any animal species (as of the fur seal, 
to take at random any one of thousands of species that 
would illustrate the point quite as well) can mean nothing 
less to a really scientific biology than the whole series of 
activities of at least one individual animal from its birth to 
its natural death. Consequently an "understanding of the 
complicated phenomena" thus presented can not be secured 

Implications of the Theories of Nerve Action 

by any amount whatever of analysis, but only through an 
endless series of never-divorced analyses and syntheses, this 
series running on through years of effort by scores of 
investigators in the field (on the Pribilof Islands, on the 
Behring Sea and far down into the North Pacific Ocean) 
and in the laboratory, by general zoologists, mammalogists, 
anatomists, embryologists, physiologists, comparative psy- 
chologists, bio-chemists and physical chemists. 

How, let one ask himself, would the resolving of a fur 
seal's behavior into reflexes as the simple elementary com- 
ponents of that behavior, contribute to an understanding of 
the annual oceanic migration of the animal or the fighting 
of the males for the females, if that analysis accomplished 
nothing beyond proving a certain measure of identity be- 
tween the reflexes of a fur seal and, for instance, those of 
an earthworm? Or how would the understanding sought 
be enhanced by carrying the analysis to a still deeper level 
to the physico-chemical level and discovering just how 
oxygen, or some particular proteid substance, participates 
in the reflex? We must never lose sight of what the biolo- 
gist's task is as regards understanding. It is to investigate 
for the purpose of understanding the facts presented by 
living beings in nature. The behavior of the fur seal as the 
animal lives its normal life is what is to be studied in the 
case chosen, and one of the most characteristic things in 
this particular behavior is a yearly journey of several thou- 
sand miles through the Pacific Ocean. That is one of the 
phenomena to be studied and understood, and no amount of 
analysis resulting in discoveries which do not apply par- 
ticularly to that particular phenomenon can be admitted 
as an explanation of that phenomenon. As compared with 
any of the other natural sciences, biology is preeminently 
the science of individuals or natural objects which though 
alike in innumerable attributes are unlike in innumerable 
other attributes, and in no aspect of organisms do the dif- 

The Unity of the Organism 

ferentials which make individuals, species, genera and so 
on come out so importantly as in behavior. 

The evidence being now overwhelming that all organic 
phenomena, including behavior, are dependent upon physico- 
chemical substances and forces, one of the most pressing ques- 
tions of procedure in biological research is that of bringing 
the older and, generally, less exact natural history aspects of 
the science into closer, more vital cooperation with its newer 
experimental and more quantitatively exact aspects. Specif- 
ically stated, work of the type long prosecuted by explor- 
ing expeditions, botanical and zoological gardens, museums, 
botanical, zoological and biological societies, and govern- 
ment biological surveys ; and that of laboratories in the 
strict modern sense, the morphological, physiological, and 
bio-chemical laboratories, must join hands more closely and 
effectively than they have heretofore to insure continued 
progress in the organic sciences. Several movements of the 
day in biology could be mentioned whose meaning, viewed 
from our standpoint, can hardly be mistaken. Perhaps the 
most conspicuous of these is that congeries of research ac- 
tivities known as ecology. In spite of frequent deprecia- 
tive comments about ecology, especially because of its in- 
definiteness as to both content and delimitation, it has the 
merit from our standpoint the very great merit of facing 
organic nature as it actually is, that is, of having for its 
subject matter the modes of life of organisms as nature 
presents them, and hence of recognizing the laboratory as 
an agency, but only as one among other agencies, for deal- 
ing with its subject. As to method, while ecology recognizes 
the indispensability of the laboratory and experimentation 
in the narrow sense, it refuses to let such experimentation 
usurp the whole of its interest and effort. 

So our study of the organism's integratedness as exem- 
plified by its activities, that is, by its behavior, and by the 
mechanism through which these activities are carried on, 

Implications of the Theories of Nerve Action 

leads to the somewhat unexpected though entirely natural 
result which may be summarily stated thus : To gain un- 
derstanding of the behavior of living beings is admitted by 
everybody to be the chief reason for investigating such ac- 
tivities. Due consideration of the nature of the activities 
and of the nature of understanding makes it certain that the 
phenomena themselves are highly integrative and integrated, 
or synthetic, and that understanding of them depends as 
much on synthetic knowledge-getting as on analytic knowl- 
edge-getting. Perception of this last truth necessitates, 
again, a sort of synthesis, or integration, of the numerous 
research agencies. 


1. Loeb ('16) 285 9. Loeb ('02) 94 

2. Loeb ('12) 70 10. Loeb ('02) 101 

3. Loeb ('16) 284 11. Schrader 177 

4. Loeb ('16) 257 12. Holmes ('06) 313 

5. Loeb ('02) 7 . 13. Holmes ('06) 315 

6. Friedlander 363 14. Luciani iii, 355 

7. Loeb ('02) 85 15. Loeb ('02) 5 

8. Loeb ('02) 91 

Chapter XXII 


Preliminary Remarks 

(a) Absolute Discrimination Between Re-flex and Psychical 

Phenomena Not Necessary 

T will be recalled that in our discussion of neural inte- 
gration we limited ourselves strictly to those manifesta- 
tions and activities of organisms which are, so far as ob- 
servation can determine, strictly reflex, that is, show no 
evidence of intelligence and volition, or even necessarily of 
instinct. What we have to do next is to consider the unity, 
the integratedness of the animal organism as manifested in 
the vast array of its activities which by universal consent 
are designated by such terms as instinctive,, emotional, vo- 
litional, conscious and intelligent. 

Be it noted that this aim will not exact of us, any more 
than did the last, a sharp delimitation between reflex or 
purely mechanical acts and psychical or conscious acts. 
Just as in the former discussion we were concerned with the 
integrative character of those acts which are indubitably 
reflex, so here our object is to study the integrative or syn- 
thetic character of those acts which are indubitably psychi- 
cal. We shall now be dealing with acts which have unques- 
tionable psychical attributes, that is, show something of 
individual plasticity and something of correspondence to 
individual needs; which are, in other words, to some degree 
individually determined to meet individual requirements 
either of external or internal imposition and intention. 


Psychic Integration 

(b) The Organism an Original Datum in All Problems of 

Psychic Life 

Another preliminary remark of high importance concerns 
the question of what, precisely, it is with which we have to 
do of what we start from and what is ever in sight, in the 
discussion. Our fidelity to the organism, living in its natural 
setting, as the foremost objective reality in this treatise, 
prevents us ab initio from being satisfied with a Body, and 
a Mind or Soul, as these have figured from time immemorial 
in discourse about the higher animals, particularly about 

If in all the world there is such a thing as objective 
truth, what we start with and have ever to deal with in 
studying psychic phenomena, just as in studying all other 
phenomena of animals, are individual objects or bodies of 
very particular construction and activity. And by no pos- 
sibility can consistent thought and statement avoid acknowl- 
edging that that vast assemblage of acts and other manifes- 
tations which are called psychical are yet only part and 
parcel of the still vaster assemblage of acts and manifesta- 
tions presented by the very same living objects, that is, by or- 
ganisms. Our occupation will be basally with an object, some 
particular organism, having innumerable attributes, which 
being classified fall rather roughly into two great groups, 
one of which we name physical or material and the other 
psychical or spiritual. For short, the physical or material 
group is called the Body, while the psychical or spiritual 
group is called the Soul or Mind. 

Our discussion, then, will never lose sight of the fact that 
the acts with which we deal are acts of the organism and not 
of any of its parts merely, whether these be conceived as 
material or psychical. No matter how far particular acts 
may be dependent upon, and so explicable by, particular 
parts, this dependence can not in reality be the whole story, 

216 The Unity of tire Organism 

for the sufficient reason that the parts are, finally, non- 
existent except as derivatives of and dependencies upon the 
organism, as our whole treatise has abundantly shown. 

Discussion of the Nervous System, the Brain, the Cerebral 
Cortex, Neurones, Reflexes, the Senses, Responses, Emo- 
tions, Consciousness, Will, Reason, and so on, as though any 
of these are now or ever have been or ever will be independent 
entities, or tilings to which the organism is subordinate, is 
from our standpoint one of the deep inadequacies and mis- 
fortunes of much biological and psychological thinking. To 
the whole attitude of the zoological naturalist, who by na- 
tive endowment and by training is imbued with the spirit 
of his motto "neglect nothing" in the study of animals, this 
liabit of the special sciences of animal life is intolerable. 

The ancient problem of the relation of "Mind" to "Body" 
is one of those problems which run on endlessly in discussion 
simply because the partisans of one theory or another never 
know exactly what they are discussing never know just 
where they start from, in what direction they are going, or 
what the end would be like if they reached it. 

Lest this statement be taken as foreshadowing both a 
right statement and a "final solution" of the problem, I 
affirm very positively that it foreshadows nothing of the sort. 
All I hope to do is to add considerably to a clear statement 
of the problem, to add a little to our comprehension of 
whither we are going, and to contribute a bit to the "final 
solution," whatever that may mean. 

The real problem of psychic integration formulates itself, 
for us, in a two-parted way: Given any particular act or 
action-system* which is unquestionably psychical, (1) how 
many and what parts of the organism are essentially in- 
volved in it? and (2) does the act or action-system bear such 

* This phrase I borrow from Jennings (Behavior of the Lower Organ- 
ixms, p. 107) and mention incidentally here that we shall find it ex- 
tremely useful later on. 

Psychic Integration 

relation to other acts or action-systems and to material 
parts of the organism as to warrant the ascription to these 
acts of causal influence on other acts and on the material 
parts ? 

(c) Provisional Classification of Psychical Facts 

A third and final introductory remark touches on the 
question of what shall be recognized as contained in the 
organism's system of psychic attributes. Following our 
regular custom of beginning with the phenomenon under con- 
templation at its fullest, most indubitable expression, we 
shall not go far amiss if we accept the time-honored trium- 
virate of feeling, will, and intellect as the most obvious sub- 
groups of highest psychic attributes ; for only a hopelessly 
sophisticated philosophy and psychology can hesitate to 
acknowledge that every full-grown, normal, civilized human 
organism, at least, is at once a sort of reservoir of feeling, 
sentiment, and emotion ; a dynamo of resolution and exe- 
cution ; and a granary of intelligence and reason. ( See, for 
example, Tlwikvng, Feeling, Doing, by E. W. Scripture.) 

Perhaps the only thing that needs saying about these 
sub-systems of mind is that our general standpoint aligns 
us squarely w r ith the tendency in present-day psychology to 
accept them for what they actually are, striving to become 
acquainted with them and to assess their importance on this 
basis. To ascertain first of all the facts on the psychic side 
of the living animal, then next to interpret, to correlate, to 
explain these facts, are cardinal principles of procedure in 
our enterprise. For one thing, as an evolutional zoologist 
of many years' practice in speculating on how animal parts 
originated (even those of almost infinite simplicity as com- 
pared with the mind of man), I am too familiar with the 
limitations and pitfalls of the genetic method to be be- 
guiled into making some one theory of the origin of mind 

The Unity of the Organism 

the corner-stone of my interpretation.* 

Furthermore, as a naturalist faithful to the mandate "neg- 
lect nothing," I am in full accord with psychology's aban- 
donment of the earlier supposition that a leading aim of 
psychology must be to prove that some one psychic province 
is all-dominant, the others being merely secondary and trib- 

Thus all forms of the theory that the psychic empire is 
at bottom Intellect, the heaven-ordained monarch of which 
is Unconditioned Ideation, are incompatible with our stand- 
point. Something of the weight and variety of authorita- 
tive sanction which are pitted against us here is indicated 
by such names as Descartes, Locke, Leibnitz, Hume, Kant, 
and Herbart of the proximate past, and Wundt, Royce, 
Howison and Bradley of the immediate past. 

And theories like those of Fichte, Schopenhauer, Hart- 
mann, Nietzsche, and divers pale, fitful present-day lights, 
which would accord to Will hegemony over the entire psychic 
realm, are still less tolerable to us. 

Feeling has won its rightful place in psychology so recently 
that there seems little danger of its pushing its claims to 
position and power beyond reason. Except perhaps in the 
form of sensationalism, or the theory that all knowledge ac- 
tually does originate in sensations, psychology of the west- 
ern world appears never to have attempted seriously the 
deification f of Feeling as it has of Reason and Will. When 

* I count it as one of the pieces of good luck in my scientific career 
that, through no merit of my own, a technical memoir of mine containing 
an elaborate theory of the origin of the vertebral column has lain in 
editorial keeping unpublished for a decade and a half. 

t It is possible, as my friend Professor G. M. Stratton suggests to me, 
that the German philosopher Fr. H. Jacobi, came nearer doing this than 
any one else. His teachings gave, however, a definite place to positive 
knowledge, so that, according to Hoffding, his faith and his knowledge 
constituted two distinct philosophies. What he wrote probably does not, 
consequently, contradict the statement in the text. Jacobi seems not to 
have exerted much influence on the main current of German philosophy 
and life. 

Psychic Integration 

Thomas Hobbes identified imagination with fancy, "orig- 
inal fancy" with sense, and "decaying sense" with memory, 
and held sense to be caused by "so many several motions of 
the matter, by which it presseth our organs diversely," and 
when he defended the general doctrine that "there is no 
conception in a man's mind which hath not at first, totally 
or by parts, been begotten upon the organs of sense," 1 he 
did indeed blaze a trail which might easily lead to an over- 
exaltation of the sensuous and emotional side of life. But 
the eminently practical character of Hobbes' undertaking 
being remembered (he was writing not for the love of specu- 
lation but to save his country from political chaos and mis- 
ery begotten as he believed from false theories and impossible 
desires and ambitions) one may expect to find in him ele- 
ments of steadiness and restraint which would make for safety 
in speculation. 

One such element is clearly seen in his opinion that he who 
is "born a man" and lives "with the use of his five senses" 
has all the native equipment necessary to realize the best in 
him both for himself and for his country. 2 The necessity 
of being "born a man" is the point to be especially noticed. 
That, with all it may imply, is on a par with the necessity 
of using the five senses. So whatever of scientific hobby- 
riding under such captions as "sensationalism," "empiric- 
ism," "associationalism," may have followed in the wake of 
Hobbes' writings, Hobbes himself, I am quite sure, was at 
heart a genuine organismalist and is entitled to high es- 
teem as one of the very first moderns to speak strongly for 
the importance of the body generally to the psychic life of 
man. Listen to this : 

"Natural sense and imagination are not subject to ab- 
surdity. Nature itself cannot err ; and as men abound in 
copiousness of language, so they become more wise, or more 
mad than ordinary." "Between true science and erroneous 
doctrines, ignorance is in the middle." 

220 The Unity of the Organism 

Having completed a reconnoissance of the field in which 
we are to work, of its expanse, its contents, and its main 
subdivisions, we are prepared to take up the task proper. 
Approaching it from our standpoint, one naturally sur- 
mises that between the organism's neural unity as manifested 
by its reflexes studied in the last chapter, and its psychical 
unity known to psychology and to be considered presently, 
a vital unity of still higher order exists. By unity I mean a 
unity so intimate, so reciprocating, so mutually constitutive 
that the term parallelism, with the meaning given it in much 
of recent psychology, is wofully inadequate for it. Such a 
unity, if it exists, must be sought by inspecting the entire 
gamut of psychic life, from the simplest responses to stim- 
uli on up through simple reflex responses, the tropisms, the 
primal affective and emotive responses, through the perceiv- 
ing, the imagining, the conceiving, the reasoning operations, 
to the very highest constructive human mental achievings. 

Likeness Between Tropistic and HigJwr Psychic Activity 

An important move, starting from the highest phase of 
rational mind, has been made toward recognizing the nature 
of this unity. This move is the more significant for our 
enterprise in that the investigator who has made it is neither 
an elementalist nor an organismalist, but an eminent sub- 
jective idealist. Josiah Royce is the student who has per- 
formed this service. Stated in a single sentence, the ad- 
vance he has made toward discovering the union consists 
in the recognition of certain fundamental resemblances be- 
tween some of the very highest operations of man's mind 
and the pure tropistic operations of lower animals. 

Royce's contribution to this subject is contained in his 
Outlines of Psychology, and nowhere else so far as I know. 
From the preface of this book I gather the following, partly 
by way of quotation and partly from obvious inference. 

Psychic Integration 

'To my mind," says Royce, "an interesting side-light has 
been shed upon the well-known controversies between the 
associationists on the one hand, and the school of Wundt 
and the partisans of 'mental activities' generally, on the 
other, by the stress that Professor Loeb has recently laid 
upon the part that what he calls 'tropisms' play in the life 
of animals of all grades." 4 

Then after telling in a few sentences what tropisms are, 
Royce continues : "Now it is especially notable that the 'tro- 
pisms' of Loeb are not, like the 'reflex actions' of the theories, 
modes of activity primarily determined by the functions of 
specific nerve-centres. Furthermore, they are more general 
and elemental in their character than are any of the acquired 
habits of an organism." (At this point Royce takes up, for 
a moment, a matter to one side of my main purpose, namely 
the problem of "self-activity" and "spontaneity" ; so I ven- 
ture to change somewhat the order and emphasis of his ar- 
gument.) "Now it has occurred to me to maintain, in sub- 
stance, that the factor in mental life which Wundt's school 
defines as 'Apperception' . . . may well be treated, from the 
purely psychological point of view, as the conscious aspect 
or accompaniment of a collection of tendencies of the type 
which Loeb has called 'tropisms.' 

Then we have : "Wundt has insisted that his 'Appercep- 
tion' is no disembodied spiritual entity. I conceive that Loeb 
has indicated to us, in the concept of the 'tropism,' how 
a power more or less directive of the course of our asso- 
ciations, and more general than is any of the tendencies 
that are due, in us, to habit, or to specific experience, can 
find its embodiment in the most elemental activities of our 

What, now, is the bearing of this idea of Royce's on the 
main theme of this chapter, the organism's unity as mani- 
fested in and influenced by its psychic life? As an initial 
step toward answering this question, the reader is asked to 

222 The Unity of the Organism 

recur to the chapter on tropistic activities and their an- 
atomical groundwork, recalling that it was the special aim 
of our discussion to show the inevitable organismal trend of 
the whole doctrine of tropisms. It should be remembered 
also that tropisms are, all of them, probably, beyond ques- 
tion adaptive in fundamental nature; i.e., they work in the 
interest of either the individual as a whole or of the species 
to which the individual belongs. The circumstance that 
under occasional more or less artificial conditions the ac- 
tivities of an animal may result in injury to it or even in its 
death, is not proof that on the whole those activities are not 
advantageous. A horse or man may make himself sick now 
and then by taking the wrong kind of food or too much 
food, but this does not prove eating to be useless on the 
whole, or non-adaptive. 

Another important thing to bear in mind about tropisms 
is their automaticity, or preferably their intrinsicality. 
They are rooted in and partake of the very essence of the or- 
ganism so much so that they manifest themselves inevitably 
when the right external and internal conditions are present, 
whether the general ends which they normally serve are at- 
tained in the particular instance or not. The flight of the 
moth toward the flame, even at the sacrifice of its life some- 
times, is a manifestation of a tendency that works, on the 
whole, for the good of the animal. That the moth follows 
the impulse even to death merely shows how tremendously 
deep-seated this type of reaction is. That the activity may 
result in injury or death in a special case is just because the 
case is special, i.e., it is a departure from the regular con- 
ditions under which the reaction-type became incorporated 
in the organization of the creature. Being always poised 
for a particular kind of action, and having a supply of en- 
ergy to execute the action, are unquestionably among the 
most distinctive attributes of animal organisms. Such or- 
ganisms are distinguished from plant organisms not only 

Psychic Integration 

by the present fact of inherent activity of the animal, but 
by their inherent preparedness for acting to meet new and 
more or less unusual situations. This action and action- 
readiness are the real meaning of the neuro-muscular system. 
All biotic organization is anticipatory in various ways, but 
animals are almost exclusively anticipatory in action. 

It is just these attributes that Royce recognizes as com- 
mon ground between certain of the highest psychic activi- 
ties of man and tropistic activities. With this overplus, 
and in some cases useless or even injurious activity (in- 
stanced by the flight of the moth toward and around the 
flame), let us now pass to the upper end of the gamut of 
animal activity for illustrations. A very few must suffice. 
The first chosen is one of exalted creativeness in art. 

From the vast domain of art a more instructive illustra- 
tion of over-wealth of self-activity can hardly be found than 
is afforded by William Shakespeare. A recent investigation 
of his works undertaken with a view to finding what they 
tell about the "native endowments of the author" and prose- 
cuted with that love for accurate, exhaustive knowledge 
which is the very soul of modern science, leads to the result 
that of these endowments "the most outstanding perhaps is 
his exuberant vitality." This characteristic of the man is 
exhibited in the "reckless volubility of almost every char- 
acter, the piling up of fancy upon fancy, of jest upon jest, 
the long embellishment of humor and foolery and horseplay 
for no other reason than the delight they afford." 7 And 
incidentally, the strict individualism of this sort of thing is 
exemplified by one of these same Shakespearian characters: 
"Come, come," says Mercutio to Benvolio, "thou art as hot 
a Jack in thy mood as any in Italy. . . . Nay, an there 
were two such, we should have none shortly, for one would 
kill the other." "What has Queen Mab to do with the ac- 
tion of the play of Romeo and Juliet? Nothing; but Mer- 
cutio mentions her, and before any one can stop him he has 

The Unity of the Organism 

poured forth fifty lines of purest fantasy. . . . Whole 
scenes," this student declares, "exist for no other reason than 
that the author's brain is teeming with situations and humor 
and infinite jest." 

Now I hear in imagination expressions of astonishment, 
rising to protest, even to ridicule, on the part of some biol- 
ogists, and to horror on the part of some literateurs, at the 
idea of suggesting that there is anything really in common 
between the two groups of phenomena here placed side by 
side the activities of a moth and of Shakespeare ! For the 
moment I do no more in reply than ask the reader to take 
cognizance of the fact that the whole training and occupa- 
tion of the naturalist consist largely in comparing all sorts 
of things, inorganic as well as organic, which to cursory 
observation seem unlike, for the purpose of finding whether 
closer and broader examination can not discover resem- 
blances and affinities which may throw light on the ever- 
insistent problem of origin and causal relationship. From 
that procedure, and that alone, initially, came the theory of 
organic evolution. It is the quintessence of the organic 
method. To him who is so instructed and disciplined that 
the recognition of likeness and kinship between, for example, 
the prothallus of the fern and the flowering plant, or between 
a horse's fore-foot and a man's hand, will receive no shock 
from the comparison. The intrinsic justification of the 
comparison will be deferred until we have a few other illus- 
trations before us. Another illustration will be taken from 
an author, J. J. Rousseau, whose activities stand about mid- 
way between art proper and science proper. 

"I felt," Rousseau says in his Confessions, "that writing 
for bread would soon have exhausted my genius, etc.," 
Again: "Nothing vigorous or great can come from a pen 
totally venal." And finally: "In a severe winter, in the 
month of February, and in the situation I have described, I 
went every day, morning, and evening, to pass a couple of 

Psychic Integration 

hours in an open alcove which was at the bottom of the 
garden. ... It was in this place, then, exposed to freezing 
cold that without being sheltered from the wind, I composed, 
in the space of three weeks, my letter to D'Alembert on 

If this sort of thing, one may note in passing, is a case 
of "struggle for existence," the existence struggled for is 
on the highest plane of psychic life and not on the plane 
of mere brute continuance. 

The only other example will be one of activity in science 
proper, i.e., "pure intellect" as far as there is such a thing. 

The case of some man devoting the best of his life to the 
working out of a great germal idea an Aristotle, a Coper- 
nicus, a Galileo, a Kant, a Darwin will serve our purpose 
best. Of these we choose the case of Darwin. Consider first 
the youth and the young man keenly alive to the flood of 
sense impressions pouring in upon him from external nature, 
and mentally "internally" "restless," as Royce would 
say, from an undefined though strong dissatisfaction with 
the stereotyped school and university curricula and modes 
of dealing with subjects. Later comes the set of environ- 
mental influences (chiefly through the naturalist Henley), 
quite incidental to his regular, prescribed environment, to 
which he responds with eagerness and effectiveness an al- 
most automatic choosing of fields of intellectual activity. 
Out of all these fragmentary and by-the-way experiences- 
"contents of consciousness" there is organized a body of 
natural knowledge, and such definiteness and promise of ten- 
dency as to justify an appointment to a post of consider- 
able responsibility and unique opportunity, that of natural- 
ist to H. H. Exploring Ship Beagle. 

During the voyage and from the new and strange con- 
tacts with nature afforded by it, there arises another state 
of "restlessness," this time concerning the origin of organic 
species, the "mystery of mysteries," as Darwin himself put 

The Unity of tlie Organism 

it. A matter deserving special notice is that the truly for- 
ward, the creative step came after, and was conditioned 
upon, a period of dissatisfaction with the prevalent teaching 
on the subject. Then the considerable time of semi-con- 
structive observation and thinking and feeling under guid- 
ance of the general surmise that species arise naturally and 
not supernaturally, as all his earlier experiences "contents 
of consciousness" -had taken for granted. And at last the 
final, for him, great conception, the hypothesis of the "strug- 
gle for existence" and "survival of the fittest" as a cause of 
the transformation of species. The suddenness and spon- 
taneousness with which this idea emerged into consciousness 
should be specially noticed. Once the merest suggestion of 
it hove in sight, the whole hypothesis formed itself, organ- 
ized itself, rapidly and completely. 

The sense in which the process may justly be called spon- 
taneous is important. Although we well know that the 
famous hypothesis was suggested by the reading of Malthus' 
work on population, we know equally well that the most es- 
sential features of the hypothesis were not contained in the 
teachings of Malthus. There was something genuinely new 
in the hypothesis. Out of the former total of experiences 
came that which did not actually exist in those experiences. 
Although the hypothesis was clearly a product of something 
which went before, it was a synthetic product in the strictest 
sense, in essentially the sense that a chemical compound is a 
synthetic product of its interacting elements, the sense that 
the most distinctive attributes of the compound can not be 
found in the elements taken separately, but only after the 
interaction has actually occurred. 

We must not fail to consider the long period of Darwin's 
strict "self-activity" in collecting evidence, pro and con, 
bearing on his hypothesis ; and the activity designed, notice, 
to ascertain whether or not there is a process going on in 
the outer world of plants and animals corresponding to the 

Psychic Integration 

process he had conceived, i.e., had pictured in his "inner 
world" of consciousness. The genuineness of the individual, 
the personal, the unique character of mental life and mental 
creation can hardly be more strikingly illustrated than by 
such cases as this of Darwin's when the conception, the hy- 
pothesis, is kept to one's self so long in order "to prove" 
whether it is "true" or not. 

Now I want to call particular attention to the indubitable 
fact that these illustrations are only extreme manifesta- 
tions of attributes which are universal in the human animal 
at least. There is no normal human known to anthropology 
which has not some measure, no matter how small, of creative 
impulse in art and in science. 

As a conclusion to this presentation of instances I must 
again insist upon one of my cardinal points : that the in- 
dividually active and creative power of the human organism 
on its psychical side is not a whit less real, less objective, less 
a natural phenomenon to the natural historian than is the 
individually creative power of physical growth and variation, 
and reflex and tropistic action. Indeed, the thorough-going, 
consistent zoological naturalist, the substance of whose 
science is largely animal behavior in all its aspects, can not 
possibly approve the effort to separate completely the two 
sorts of creation. 

First Move Toward Showing the Organismal Character of 

the Higher Psychic Life 

Now for the further scrutiny of such psychical facts as 
those typified by the examples presented, for the pur- 
pose of seeing what has been done and may yet be done to- 
ward bringing them into accord with the organismal con- 
ception, the pole star of all our previous discussions. This 
examination will begin, as others have begun, by showing 
how elementalistic attempts to interpret organic phenom- 

The Unity of the Organism 

cna soon reveal their inadequacy and finally break down as 
the efforts come to face the increasing complexity which 
progress of objective research always finds in such phenom- 

Associationist Psychology a Special Case of Element alist 


In the particular psychical realm we are now to examine, 
elementalist theory has appeared most prominently as what 
is called Associationism. This flourished first in England as 
the school of English Associationists, David Hartley, near 
the middle of the eighteenth century, being usually consid- 
ered its founder. Psychologists of this group hold that 
ideas, which for them appear to be identical with sensations, 
are the "ultimate elements" of psychic phenomena. "Ac- 
cording to this theory, rigidly carried out, all genesis of new 
products is due to the combination of pre-existing ele- 
ments." Even the passions, according to Hartley "must 
be aggregates of the ideas, or traces of the sensible pleas- 
ures and pains ; which ideas make up, by their number and 
mutual influence upon one another, for the faintness and 
transitory nature of each singly taken." The "piling up 
of fancy upon fancy, of jest upon jest, the long embel- 
lishment of humor and foolery and horseplay" which Pro- 
fessor Manly shows characterize many of the Shakespearian 
plays, would be explained, according to this kind of psy- 
chology, not really by the author Shakespeare but by the 
"aggregation," in some way, within him of ideas. 

And, similarly, the works which in popular language are 
said to be by a Darwin, a Humboldt, a Copernicus, an Aris- 
totle, are in reality not by but merely m these men. The men 
were only the places of aggregation of the elements the 
ultimates by which the teaching on the origin of species, 
on the general character of the earth, on the solar system, 

Psychic Integration 

on the deeper meaning of external nature, were produced. 
Again the old story with which we have become familiar: not 
the organism, but elements of, or perhaps merely in it, are 
the causal explanation of whatever occurrences are associ- 
ated with the organism. It is, I think, safe to assume that 
both the merits and the demerits of associationist psychol- 
ogy have been made patent enough, at least to English- 
speaking students, by the writings of James and others. 

If only the doctrine of "association of ideas" can be satis- 
fied to do what it is really able to do, and not insist upon 
trying to do what it can not do, its usefulness is great and 
its permanence in psychology assured. 

As indicated above, the "huge error," as James expresses 
it, by which the "whole historic doctrine of psychological 
association is tainted" ' is only another miscarriage of the 
elementalist mode of reasoning, and so is subject to the 
general type of criticism which the reader has met in every 
chapter in this book. 

In order to divest the criticism as much as possible of 
personal flavor I shall make large use of James' language. 
"All these writers," says James, referring to Hobbes, Hume, 
Priestley, Hodgson and the later English associationists, 
"hold more or less explicitly to the notion of atomistic 'ideas' 
which occur. In Germany, the same mythological suppo- 
sition has been more radically grasped, and carried out to 
a still more logical, if more repulsive, extreme, by Herbart 
and his followers, who until recently may be said to have 
reigned supreme in their native country." 

Now the objection to the doctrine of "atomistic ideas" 
does not so much concern the conception of ideas as atoms 
as the nature attributed to these atoms, namely in assum- 
ing them to be immutable, and sufficient in their isolate 
capacities to account for the thought and other products 
arising from their "association." The following two quota- 
tions illustrate the form this criticism, the essence of which 

230 The Unity of the Organism 

is now very familiar to us as biologists, takes when it ap- 
pears in garments of a psychologist's making. The "huge 
error" of the association doctrine, mentioned above, James 
explains, is "that of the construction of our thoughts out 
of the compounding of themselves together of immutable 
and incessantly recurring 'simple ideas.' 

If there be any doubt as to the meaning of this surpris- 
ingly un-James-like wording, there certainly can not be as 
to the following: "For Herbart each idea is a permanently 
existing entity, the entrance whereof into consciousness is 
but an accidental determination of its being. So far as it 
succeeds in occupying the theatre of consciousness, it crowds 
out another idea previously there. . . . The ingenuity with 
which most special cases of association are formulated in 
this mechanical language of struggle and inhibition, is great, 
and surpasses in analytic thoroughness anything that has 
been done by the British school. This, how r ever, is a doubt- 
ful merit, in a case where the elements dealt with are arti- 
ficial ; and I must confess that to my mind there is something 
almost hideous in the glib Herbartian jargon about Vorstel- 
lungsmassen and their Hemmungen and Hemmungssummen, 
and striken and erheben and schweben, and Verschmelzungen 
and Complexionen." ll 

The long and short of the "huge error" of associationist 
psychology is that ideas are no such independent, immutable, 
simple entities as the doctrine supposes ; that in their origin 
and in all they are, and all they do, and all that comes forth 
from their association, they are in some sort and measure 
dependent upon what? Something. Search after this 
something has been a large motive of more recent psycho- 
logical inquiry. 

One way of supplementing and rectifying associationist 
doctrines is to epitomize the shortcoming of these doctrines 
in the statement that they recognize only the objective side 
of the association process, whereas the subjective side is 

Psychic Integration 

equally important. Thus Pillsbury : "It was a neglect of 
the subjective conditions and the insistence upon the objec- 
tive side of the problem that has led the English Associa- 
tional School into disrepute. The explanations that they 
gave were true as far as they went, but their incompleteness 
vitiated the conclusions as soon as they laid claim to uni- 
versality." And the author then shows, convincingly 
enough, how two sets of subjective factors, attention and 
choice, play a large and important role in determining the 
"associative train." And further, "A complete explanation 
of association demands that both sets of factors [objective 
and subjective] be taken into account; to omit either is to 
fail in the solution of the problem." 

Preliminary Examination of Objective and Subjective 

There is undoubtedly a real gain in having proved, as 
Pillsbury and others surely have, that a "side" other than 
the objective in association does exist. It is highly ad- 
vantageous, also, to have learned enough about this other 
"side" to make the term subjective an appropriate name 
for it. But any one coming to a study of the associational 
activities of the organism's psychic life as we have, namely 
as naturalists, can not avoid, if true to his traditions and 
methods, wanting to know how these two "sides," the objec- 
tive and the subjective, go together what the nature is of 
their relation. For the very fact that they are two sides 
of one thing is to the naturalist prima facie evidence that 
they are in vital organic connection. Even the two sides of 
an inanimate thing, like the earth with its two hemispheres, 
have a relation to each other too important for the earth 
sciences to ignore or even to put off as merely "parallel." 
But when it comes to an entity like a live animal, the quin- 
tessence of which is organization, the question of how two 
of its "sides" so important as its objective and its subjec- 

The Unity of the Organism 

tive are related, becomes most fundamental, especially when 
a subject like that of psychical association is up for con- 

It is, then, fundamental to our enterprise to find out all 
we can about the connection between the objective and sub- 
jective aspects or sides of the organism. It would be folly 
to expect results of any value from an effort of this sort 
without having first given attention to the nature of each 
of the sides. Now the objective "side" comes to much the 
same thing as the physical or material organism as we are 
conceiving the "sides." But since this has been the sub- 
ject of our whole treatise up to the last two chapters, and 
even of the greater part of these, we are already possessed 
of enough understanding of this "side" for our present 

As to the subjective "side" the case is different. Into 
its nature, its makeup and activities, we have looked very 
little only in a bird's-eye-view fashion thus far in the 
present chapter, and into its marginal or transitional zone 
in the preceding two chapters. We are, consequently, 
obliged to penetrate further into the subjective realm itself 
before the main problem, that of the relation between the 
sides, is attacked. 

The Essence of Wundtian Apperception 

This carries us back to the point at which we brought 
Royce's suggestion of a relation between Loeb's tropism 
theory and Wundt's apperception theory into the discus- 
sion, the return to this point being for the purpose of using 
Wundt's conception to induct us further into the nature of 
mental life. The importance of examining Wundt's con- 
ception is two-fold. In the first place, we want to know 
whether or not it is genuinely descriptive of man's highest 
psychical life. If it is, nothing can stand in the way of its 

Psychic Integration 233 

acceptance, so far, by the anthropological zoologist. But in 
the second place we must know whether or not it carries, as 
some critics believe it does, transcendental or supernatural- 
istic implications. If this charge be true it is of course to 
this extent unsanctionable from our standpoint. 

Wundt's most concise characterization of apperception 
which I have found is, "The process by which any content 
of consciousness is brought to clear comprehension we call 

A content of consciousness is any definable experience we 
may have. All consciousness whatever is consciousness of 
something or other. This "something or other," no matter 
what, nor whether regarded as a whole or in part, is a con- 
tent of consciousness, according to my understanding. 

What is most distinctive about Wundt's characterization 
may be regarded as centering around the word clear. When 
a particular content gets itself into the lime-light of con- 
sciousness when it becomes the center of attention the 
process by which it does so is apperception. On the other 
hand, the process by which contents, though brought into 
consciousness, come only into its outer zones or edges, and 
do not monopolize attention, is perception. Though this 
getting of a content into clearness in consciousness, this 
monopolizing of attention, may take place passively or 
actively so far as the mind as a whole is concerned, the ac- 
tive way seems to be the more distinctive and important, at 
least for mental life as a whole. 

It is apparently this positive activity of apperception, 
directed toward making particular contents of conscious- 
ness clear, which has brought criticism upon Wundt's con- 
ception. "Wundt talks," says Pillsbury, "almost as if 
there were a faculty or force of apperception, something 
behind and superior to consciousness, which brings about 
the change in clearness of the impressions. There is in 
the brain a definite centre of apperception, and in conscious- 

234 The Unity of the Organism 

ness a force very closely related to will, that in and of itself 
chooses certain ideas for elevation to the high places of 
consciousness, and equally arbitrarily rejects others." And 
then follows this in Pillsbury's criticism, which brings out 
unmistakably its real purport: "It is very much like the 
self-conscious unity of apperception of Kant, which gives 
the final form and order to the various disconnected elements 
of the mind, and is in so far something inexplicable, a factor 
in experience that must be assumed without any further 
discussion of its nature, origin, or laws of action." 15 A 
suspicion, obviously, that the transcendentalism of Kant 
broods over Wundt's theory. As to the justification of this 
suspicion we need not be concerned here. Enough for 
us at this point to recognize that from the standpoint of 
description as natural history practices the art, or aims to 
practice it, Wundt's account of the way the mind works in 
a vast range of its activities seems true, and as far as it 
goes is satisfactory. 

Not only the matter of clearness of the contents of con- 
sciousness, but their makeup as well is important. Al- 
though "psychical elements" figure largely in Wundt's sys- 
tem, one finds no intimation that the whole mind and its 
contents can be "explained" by reducing them to "ultimate 
elements" after the familiar manner of elementalist explana- 
tion. "All the contents of psychical experience," Wundt 
says, "are of a composite character." And it follows from 
this that "psychical elements, or the absolutely simple and 
irreducible components of psychical phenomena, are the 
products of analysis and abstraction." 

The two words, "analysis" and "abstraction," need par- 
ticular consideration. The psychical elements found by 
analysis do not exist, as such, in nature. Analysis, in this 
case, is logical or thought-analysis, and not objective analy- 
sis. We should do well to recall what was said in the dis- 
cussion of reflexes, namely that the "simple reflex," though 

Psychic Integration 235 

legitimate and useful as an aid to interpreting the phenom- 
ena of reflex nerve action, has no actual existence in nature. 
It, like the "psychical element," is an abstraction. This is 
only another way of saying that psychical elements are 
what they are because they are parts of the mind as a whole, 
just as we have seen over and over again physical elements 
of the body are what they are because they are parts of the 
body. "The specific character of a given psychical process 
depends for the most part not on the nature of its elements 
so much as on their union into a composite psychical com- 
pound. Thus, the idea of an extended body or a rhythm, are 
all specific forms of psychical experience. But their charac- 
ter as such is as little determined by their sensational and af- 
fective elements as are the chemical properties of a compound 
body by the properties of its chemical elements. Specific char- 
acter and elementary nature of psychical processes are, ac- 
cordingly, two entirely different concepts." 

We must not miss an essential point in this, that since 
with psychical elements just as with chemical elements we 
never know exactly what or how much each particular ele- 
ment contributes to the compound, we are obliged to con- 
ceive the attributes of the compounds as pertaining to the 
elements collectively even before the compounding has been 

So it comes about that because all contents of conscious- 
ness are fundamentally composite, but are also resolvable into 
components of various grades of complexity, synthesis and 
analysis have a prominent place in the Wundtian system. 
Of these, synthesis is the more definitive and fundamental, 
since it enters into the very nature of consciousness itself. 
Consciousness is, according to Wundt, the :< inter-connec- 
tion of psychical compounds." "It is the name for the gen- 
eral synthesis of psychical processes, in which synthesis the 
single compounds are marked off as more intimate combina- 
tions." 18 The meaning of this is made clearer by the state- 

36 The Unity of the Organism 

ment that unconscious states like deep sleep, faint, and so 
on, are the interruption of these interconnections. 

Remarks On Analysis and Synthesis 

This brings us to where we can see the important distinc- 
tion between an aggregation and a synthesis in a psychical 
sense particularly and likewise between a fragmentation 
and an analysis. No mere aggregation,* as of ideas or emo- 
tions, would make consciousness. Only a synthesis of con- 
stituents can do that. And, contrariwise, while the mere 
severance of the synthesized components produces uncon- 
sciousness, an analysis of them results, not in unconscious- 
ness, but in a consciousness of the constituent parts of the 
contents of consciousness. The essence of consciousness is 
unitariness integratedness, in our general terminology as 
regards the contents of an individual organism's psychical 
nature, so that whatever analytical processes the mind per- 
forms must move within the bounds of its own unitariness 
or integratedness. Were we to conceive the analytic opera- 
tions of the mind to exceed or even quite to equal its synthetic 
operations, we should have to conceive it as utterly negating 
consciousness, i.e., as destroying itself. A man could ana- 
lyze his own mind in an elementalist sense only by suiciding. 
In other words, he could never do it, simply because he 
would have killed himself by the very process of analyzing 
before he had completed his job. 

These remarks on the distinction between synthesis and 
aggregation, and between analysis and fragmentation, are 
not quite what Wundt says. They go somewhat beyond his 
actual expression, but are legitimate inferences, I am quite 
sure, from his discussion as a whole. And they help us toward 
what we want to accomplish, namely to discover still more 

* Recall our previous remarks on this subject, e.g., pp. 182 and 268, 
and also the quotation from Hartley, p. 228. 

Psychic Integration 237 

than Royce discovered about the relation between appercep- 
tive processes, as Wundt conceives them, and the processes 
known as tropisms. 

Anticipating our results, and stating them in the most 
general terms possible, we may say that the "apperceptive 
synthesis" of Wundt, and what may be called tropistic syn- 
thesis, have a common ground in the kind of synthesis which 
is the very essence of that kind of organization to which the 
term life is applied. To be alive is to be an organic in- 
dividual ; to be an organic individual is to be an indi- 
vidual that perpetually synthesizes itself from substances 
extraneous to itself (food, in the narrower sense, and 
oxygen) ; and to be a psychically endowed individual 
is to be an individual which in addition to synthesizing 
a physical nature from the substances mentioned, synthesizes 
a psychical nature from physical and chemical contacts and 
interactions between the individual and the external world, 
the physical contacts being called stimuli. Viewing the 
matter thus, it is seen to be highly that in its ulti- 
mate essences the dependence of the psychical nature of the 
organism on stimuli is connected, directly and inseparably, 
with the dependence of its material nature on material nu- 

We should, I think, be surprised were a demonstration 
to be produced that psychic life has as little connection with 
metabolic processes as the text-books of psychology would 
lead one to suppose. Every modern psychologist, like every 
modern biologist, accepts, unquestioningly I presume, the 
conception that in some way the psychic life is no less de- 
pendent on the nutritive substances and processes than is 
the physical life. Yet that "some way" appears to be gen- 
erally regarded as so remote and obscure as to be beyond the 
reach of profitable treatment by psychological science, judg- 
ing from the considerable number of standard text-books 
which I have consulted on the point. In only one of these 

238 The Unity of the Organism 

(Elements of Physiological Psychology, by Ladd and Wood- 
ward) do I find the word "metabolism" in the index. 

Our task may then be restated as that of making out 
more fully and clearly than Royce did the connection be- 
tween apperception and tropisms, which is involved presum- 
ably in the whole problem of organic synthesis from its 
highest manifestations in psychic synthesis to its lowest 
manifestation as metabolic synthesis ; of bringing to more 
specificity the general statement made above. But such 
statements are altogether too sweeping and abstract to satis- 
fy scientific description and explanation in our day. A 
chapter must now consequently be devoted to making them 
more definite. 


1. Hobbes 15 10. James ('90) I, 553 

2. Hobbes 21 11. James ('90) I, 603 

3. Hobbes 25 12. Pillsbury 106 

4. Royce ix 13. Pillsbury 112 

5. Royce x 14. Wundt 229 

6. Royce xi 15. Pillsbury 270 

7. Manly 3 16. Wundt 32 

8. Manly 4 17. Wundt 33 

9. Baldwin I, 80 18. Wundt 223 

Chapter XXIII 



A Still Closer Look at the Organismal Nature of Tropisms 

T T AVING selected tropisms as a strategic point in our 
program of search for the vital connection, if such 
exists, between the physical and the psychical, we must turn 
again to this subject. Our previous treatment of the tro- 
pism theory brought out the essential organismal character 
of the type of activity to which the term tropism has been 
applied. The result of that treatment might be epitomized 
by saying that in so far as the theory rests on accurate 
and adequate description, it is genuinely organismal and 
genuinely sound, but in so far as it rests on causal ex- 
planation that is elementalistic in spirit and expression it is 
genuinely unsound. Our present aim will be furthered by 
illustrating this epitomized stricture on the theory in a little 
different way from which we objectified our criticism in the 
earlier treatment. 

Every one familiar with current explanatory discussion 
of tropisms must have noticed the large and free manner in 
which the word substances is made use of in the explanations. 
Thus, to illustrate: The larvae of certain butterflies emerge 
from their winter nests under the influence of the warm 
spring sunshine, crawl to the tips of the branches of some 
shrub or tree, eat the buds and tender leaves there; then, 
after feeding to satiety "turn tail" and crawl down the 


240 The Unity of the Organism 

branches. This rather complex and, to the insects, highly 
useful performance Loeb and others have proved to consist 
of a series of reflexes so interconnected as to come under 
the tropistic type of activity. And Loeb, e.g., in the chap- 
ter, On the Theory of Animal Instincts (Physiology of the 
Brain) uses the case to good effect in support of his 
contention that the traditional instinct-and-nerve-center ex- 
planation of such phenomena is utterly inadequate. 

So far, good. As to the straightforward presentation of 
facts, Loeb's position seems unassailable. But what about 
the causal explanation of the facts? What, exactly, is it 
that sends the larvae up the branches? What causes the 
eating activities? What makes the creatures then turn 
about and finally sends them down the branches? That 
several environmental factors, the warm weather, the 
sunlight, the character of the plant buds and leaves, and so 
on, are involved is brought out clearly enough. But what 
about the factors pertaining to the larvae themselves? The 
body-shape, the skin, the sense organs, the muscles are, as 
was emphasized in the previous discussion of tropisms, freely 
recognized after a fashion by the tropism theory. But 
deeper still than these what? Chemical substances "ac- 
cording to requirement," in the language of the cook books. 
Until the caterpillars have taken food they are positively 
"heliotropic," that is, literally, are induced by sunlight to 
move toward the sun, after the higher spring temperature 
has caused chemical changes in their bodies essential to 
such movement. But by eating to satiety the chemical 
changes essential to the positive heliotropism are inhibited 
and a negatively heliotropic state comes on. "We can im- 
agine," writes Loeb, "that the taking up of food leads to 
the destruction of the substances in the skin of the animal 
which are sensitive to light, upon which substances the helio- 
tropism depends, or that through the consumption of food 
the action of these substances is indirectly prevented." * In 

Organic Connection Between Physical and Psychical 

the total scheme, then, actual and imagined, various "sub- 
stances" are indispensable. 

That the imaginary constituents constitute a very im- 
portant part of the explanation is obvious. This fact is, 
however, not specially objectionable. It is not if its true 
character is never forgotten. But here comes the point I 
wish to make focal just now. If imagination is to be given 
a place at all in the argument it must have a larger place 
than Loeb has accorded it. Otherwise the teachings of 
evolution, i.e., the genetic continuity in biology, are tacitly 
repudiated. Attention has previously been called, especially 
in the chapter on the organism and its chemistry, to the deep 
current of virtual anti-geneticism which runs through physi- 
ology generally, and particularly through bio-chemistry. 
Undoubtedly we can imagine "substances" produced and de- 
stroyed in such a complex of activities as that described, to 
meet exactly the needs of the larva ; but can we legitimately 
imagine them to be so produced and so destroyed by any 
other means than just by the particular animals in question, 
that is, by the organisms? Various of our discussions, but 
particularly those in which the specificity of protoplasm 
have been dwelt upon, constitute a decisive negative answer 
to this question. Xo causal explanation of the requisite 
"substances" imagined can stop short of the organism, alive 
and normal, as an essential and "causal factor" in the phe- 
nomena presented. Causal explanation of tropisms which 
aims to reach a physico-chemical basis is really organismal 
as well as are tropisms seen through the medium of pure 

The Automatic and Anticipatory Character of Tropisms and 

Other Re^exes 

Nor should the reader fail to note the intrinsicality, the 
adaptiveness, and the anticipatoriness, of tropisms as illus- 

The Unity of the Organism 

trated in this example. Conscious will and choice seemingly 
do not come into the operation at all. Given the right con- 
ditions, internal and external, the caterpillar goes through 
the concatenation of operations necessary for its existence, 
willy-nilly. Moreover, the actions initiated by the warm 
weather, the larvae being yet down at the base of the shrubs 
or branches, have in organic prospect, as one might say, a 
supply of food peculiar to the species. And this supply, 
be it remarked, is several inches at least, and several min- 
utes at least, away from the larva at the beginning of its 
round of activities. Its future, even more obviously than 
its present, existence is involved in the acts. Anticipatori- 
ness is perhaps the most conspicuous attribute of the adap- 
tiveness of such activities. C. Lloyd Morgan has well ex- 
pressed the truth that one of the most important lessons 
to be learned from the study of animal behavior "in its or- 
ganic aspect" is the fact that "living cells may react to 
stimuli in a manner which we perceive to be subservient to a 
biological end, and yet react without conscious purpose 
that is, automatically." 2 

But from our examination of the cell-theory we conclude 
that "living cells" in this statement ought to read "living 

So much by way of further preparation, in the reflex and 
tropistic phases of animal life, for our search after a vital 
connection between the physical and the psychical. It will 
now be advantageous to return to that supremely important 
aspect of human psychic life already examined somewhat, 
namely that of Wundtian apperception. 

A Still Closer Look at the Likeness Between Higher Ra- 
tional Life and Tropisms 

As Royce's statement of the objections to the concep- 
tion of Wundt contains several points that will be useful to 

Organic Connection Between Physical and Psychical 

us, we reproduce more of his sentences. "It has been ob- 
jected to the partisans of Wundt that the term 'appercep- 
tion,' as thus used, seems to signify a factor in mental life 
which can be explained neither in terms of what we have 
called sensitiveness, nor in terms of the law of habit. It 
has also been objected that the conception of a conscious 
process, engaged in influencing its own states, is a concep- 
tion which confuses together metaphysical and psycholog- 
ical motives. The psychologist, engaged as he is, not in 
studying how Reason forms the world, but in observing and 
reducing to rule the mere phenomena of human mental life as 
they occur, is not interested, it has been asserted, in a power 
whose influence upon mental phenomena seems to be of so 
ambiguous a character as is that which the Wundtian 'ap- 
perception' possesses." 3 

Again : "This is the place," Royce writes, "neither to 
expound nor estimate Wundt's theory. But it does here 
concern us to point out that what occurs in mind whenever 
we are actively attentive is attended with a feelmg of rest- 
lessness, which makes us dissatisfied with all those associa- 
tive processes that do not tend to further our current in- 
tellectual interests. On the other hand, the cerebral proc- 
esses that accompany active attention are certainly such as 
tend to inhibit many associative processes that would, if 
free, hinder our current intellectual interests." Mean- 
while, "our active attention itself is always the expression 
of interests which possess the same elemental character that 
we have all along been illustrating in the foregoing para- 
graphs. The attentive inventor is eager about the beau- 
tiful things that he thinks of while he is trying to invent. 
The attentive hostess is eager about social success. The 
attentive caged animal is eager about whatever suggests a 
way of escape." 

The discussion from which these sentences are taken is 
contained in a chapter near the end of the book, entitled 

244 The Unity of the Organism 

The Conditions of Mental Initiative, and in order that the 
reader may get the full force of what Royce is talking 
about, he is earnestly recommended to read the entire chap- 
ter. Only thus can the "foregoing paragraphs" mentioned 
be adequately appraised. But we must try, in our own 
waj r , to get the essence of the matter. Royce's presenta- 
tion is his way of insisting upon the facts of psychical life 
and activity, high and low, which have given rise to the 
Wundtian conception of apperception, these facts being 
the indubitably initiatory, directive and selective qualities 
of mind in all its grades. Furthermore, Royce dwells on 
the homogeneity, as one may express it, of this intrinsicality 
of mental life its initiative, its persistence, and its selectiv- 
ity with the individual or fluctuating variations which 
have played so large a part in theorizing about organic 
evolution and heredity during the Darwinian era of biology. 
And he goes back still further in good modern biological 
fashion, and connects these variations with organic growth 
itself, thus calling attention to the fact that variations of 
this particular sort can not be referred to environmental 

At this point we may stop, as biologists, to supplement 
Royce's argument by pointing out that variations of the 
sort indicated, are referable to environmental influence only 
in the sense that growth is so referable. An organism's 
securing and taking in of its nutritive substances are un- 
doubtedly a kind of response to contact with its environ- 
ment, and in that broad sense growth may be said to be 
due to environmental influence. If the organism had no 
nourishment, if it received no environmental influence of this 
kind, it certainly would not grow. At the same time, since 
the organism manages somehow to build a great variety of 
tissues and organs out of one and the same supply of nour- 
ishment; that is in response to one and the same "environ- 
mental influence" (as we are agreeing to use the phrase 

Organic Connection Between Physical and Psychical 

here) there is no course open but to recognize that the or- 
ganism is a very important, because indispensable, factor in 
its own growth and differentiation. "Self differentiation," 
so far as the whole organism is concerned, is a fact than 
which no other in the whole domain of biology is better es- 
tablished. Indeed, self differentiation is really a special 
form of self growth and surely no one would contend that 
environmental influence is more than an essential factor in 
the growth of an organism. To hold it to be a complete 
explanation of the phenomenon would be too manifestly ab- 
surd to receive serious consideration. It would be to con- 
tend, in effect, that one of the processes of the organism 
(its growth) is more than all the processes of the whole or- 
ganism. But since most if not all variation depends, either 
directly or indirectly, upon growth, what is more natural 
than that the living, growing organism should display much 
self variation? 

That such variations are among the most common phe- 
nomena presented by organic beings, there is no shadow of 
doubt to any one who views the problem broadly and crit- 
ically, and with no domineering preconceptions as to what 
ought to be and ought not to be ; who, in other words, views 
the organic world as a natural historian, guided by the 
mandate "neglect nothing," instead of as a physicist in the 
mathematico-laboratory sense, guided by the mandate "neg- 
lect everything which can not be made to conform to gen- 
eral mathematically statable law." 

These remarks about the relation of mental activity to 
growth, differentiation and variation of the organism, and 
to environmental influence would apply throughout, muta- 
tis mutandis, to tropistic activity. 

246 The Unity of the Organism 

A Still Closer Description of the Subrational Mowty of 

Psychic Life 

And this brings us to where our final return may be made 
for purely descriptive and comparative purposes, to the 
subrational moiety of psychic life, the purpose of the re- 
turn this time being to characterize this moiety as faith- 
fully but as briefly as possible on the basis of the total re- 
sults of researches in the field up to the present time. So 
bulky and varied are these results that to examine them 
exhaustively is hardly possible for any one person even 
though he be a specialist in the field. Much less possible is 
it, then, for a general zoologist to make such an examina- 
tion. Nevertheless it is, I believe, possible to give an 
epitome of the present state of knowledge that shall be true 
in all fundamental respects and highly significant for our 

Remarks on the Classes of Subrational Life 

In giving this epitome we shall not try to maintain a sharp 
distinction between reflexes, whether of the tropistic or any 
other type, and instincts. To begin with, as always, when 
a large and intensively cultivated domain of science is en- 
tered for the purpose of extracting from it its most certain 
major results, we may take it for granted that the ex- 
tremists touching any portion of the field aver which di- 
vergence is wide and warm, are unsafe guides for the gen- 
eral student. Thus, the student who enters the realm of 
animal behavior for such a purpose as that for which we 
are now entering it soon sees that those specialists who find 
nothing but tropisms, and these of the most uncompro- 
mising sort, in the activities of much of the animal king- 
dom, are not the ones to whose guidance he can entrust 

Organic Connection Between Physical and Psychical 

himself, no matter how voluminous, and perhaps excellent 
in quality, their experimental researches may be. 

For instance, such a view as that of Bohn's, according to 
which the word instinct ought to be eliminated from the 
terminology of science "as a legacy of the past, the middle 
ages, the theologians and the metaphysicians," is so obvi- 
ously unjustifiable to any well-informed zoologist as to 
make him suspicious of such a writer all along the line, 
especially wherever his judgment and scientific poise are im- 

This question of the reality of instincts I use to illus- 
trate the peril to the general student of the unpoised spe- 
cialist, because it is germane to the present discussion. In 
general zoology the type of animal behavior to which the 
term instinctive is applied is not less conspicuous and real, 
to say the least, than is the type described as tropistic. 
For an experimentalist to come out of his laboratory and 
tell a broadly experienced entomologist or ornithologist, for 
example, that the familiar achievements of young insects 
of many species, and of numerous young birds should not 
be called instinctive because (as the experimenter asserts) 
they are reducible to the tropistic or perchance the simple 
reflex type of reaction, may justly be characterized as sci- 
entific impertinence. It is as though an embryologist, hav- 
ing discovered that a bird's wing is the genetic counterpart 
of a salamander's forelimb, should instruct the ornitholo- 
gist that it is wrong for him to call the bird's wing a wing, 
because the member may be reduced to a lower type of limb. 
Unquestionably the experimental specialist often pro- 
duces results which necessitate changes in the general zo- 
ologist's conceptions and nomenclature. But it is not his 
province to take into his own hands the revision of the 
fundamental terms of zoology. Any one moderately in- 
structed in the history of zoology knows that "instinct'' is 
a hardly less well-grounded zoological term than "birth" or 

2-48 The Unity of the Organism 

"intelligence," or many another indispensable term. 

Our inquiry is not as to whether there are such things as 
instincts, but how they operate and what they signify for 
the animals possessing them. Perhaps of highest interest 
to us is the fact that innumerable instincts, if indeed not all, 
are as indubitably hereditary as are any animal endowments 
whatever. This comes out especially convincingly in those 
numberless cases where the instinctive operations develop 
strictly part passu with the anatomical development of the 
young, there being absolutely no opportunity for them to 
learn, even subconsciously. 

Take as an example the crustacean Amphithoe longimana, 
in which Holmes compared in detail the activities of the 
newly hatched young with those of the adult. "Amphithoe 
lives in tubular nests which are usually lodged among sea 
weed. The nests are somewhat longer than the animal, 
and are spun of a web-like material into which bits of sea 
weed are often incorporated which help to conceal the oc- 
cupant. In its nest Amphithoe lies in wait for prey, ready 
to dart out upon any small creature which touches the ends 
of its long antennae. 

"The activities of the adult Amphithoe, with the exc\p- 
tion of those concerned in reproduction, are almost ex- 
actly paralleled by those of the young. I have taken the 
eggs from the maternal brood pouch shortly before hatch- 
ing and kept them isolated in individual dishes. For some 
time after emerging from the egg the young were weak and 
had imperfect control of their movements, which were jerky 
and irregular. Soon the minute creatures would crawl and 
swim much like the adults, and the next day they began 
constructing nests which were the same shape as those formed 
by their parents." Then comes a part of the description 
to which the reader's special attention is called because it 
brings out, partly by implication, a richness of detail in be- 
havior which defies full expression, and which every care- 

Organic Connection Between Physical and Psychical 249 

fully observing zoologist knows to be characteristic of the 
activities of nearly all animals. Especial attention is in- 
vited to this, because this elusive wealth of behavior is 
usually overlooked by the cursory observer on the one hand 
and by the experimentalist on the other. "The attitudes in 
the nest," Holmes writes, "the waving of the antennae, the 
beating of the swimmerets, the restless movements of the 
legs and mouth-parts, springing after food, belligerency 
toward passers by, the little unobtrusive signs of timidity, 
the reversal of position in the nest on the approach of 
danger and the general behavior outside of the nest, were, 
on the next day after hatching, almost exactly the same as 
in older individuals. The only differences in behavior were 
due to the feebleness of the young and their imperfect con- 
trol of their movements." One never reads a description 
like this by a typical experimentalist, especially if he be a 
pure tropist, or by a meagerly trained zoologist ! Then the 
final statement : "The young are hatched with all the in- 
stincts necessary fully to equip them for the business of life. 
No experience is necessary to teach them what is advan- 
tageous for them to do." 6 


'The impossibility should be noticed of drawing a sharp 
line in this description between instinctive and purely reflex 
acts. "Reversal of position in the nest on approach of 
danger" is clearly instinctive. But "beating of swimmer- 
ets," and especially the "restless movements of the legs" 
are these instinctive or wholly reflex? Probably they are 
reflex, though the leg movements may well be partly in- 
stinctive. A whole volume of examples as unquestionable 
as this could be compiled, and all groups of animals from 
mammals down to worms at least would be represented. 

250 The Unity of the Organism 

Four Certainties About the Adaptiveness of Subrational 

Psychic Activities 

Concerning the purposefulness or adaptiveness of activi- 
ties of this general type, I think four things may be re- 
garded as absolutely certain. 

Generally Useful to Individual and to Species 

First, a vast majority of them are recognizably contribu- 
tory to the perpetuity of both the individual in its normal 
life, and of the species. But for them neither individual 
nor species would continue to exist. This is so obvious that 
further remark upon it is unnecessary. 

Many Useful to Species Primarily 

Second, in a large number of instances particular acts 
by particular individuals are in the interest of the species 
primarily and of the individuals only secondarily or not at 
all. This is shown most conclusively in cases like that of 
several species of salmon, where the individual normally goes 
through activities which secure the continuance of the spe- 
cies but which end in the death of the individual. A large 
and varied number of cases of this type occur, especially 
among insects. But the supremacy of species over individ- 
ual needs appears under various other forms. Thus almost 
certainly such tropistic activities as that of the moth going 
to its death or injury in the flame is of this sort. This 
case may be stated in general terms thus: Owing to lack 
of any ability on the part of an individual to modify its in- 
herited mode of action to meet a special situation, it acts in 
the old way even though the new situation, while in gen- 
eral like the old, yet differs from it enough to make it peril- 
ous to the individual if it acts unmodifiedly in the old racial 

Organic Connection Between Physical and Psychical 251 

way. The preeminently racial utility and hereditary char- 
acter of instincts is certainly one of the most interesting 
things about them for the present discussion. 

Variability of Sub-rational Activities 

The third certainty about reflex and instinctive activities 
is that they are by no means so stereotyped and invariable 
as older cursory observation or as much theorizing, espe- 
cially about tropisms and instincts, has held them to be. 
Darwin, in the notable chapter on Instinct in The Origin of 
Species, was the first to attack seriously the notion of such 
invariability in dealing with instincts. He undertook to 
show that the instinctive type of activity is subject to vari- 
ation just as are all other aspects of animal life. 

A telling set of recent investigations under this head is 
by the Peckhams. That on the solitary wasp, Ammophila 
urnaria, is particularly to the point because the earlier 
writers had used its habits of paralyzing caterpillars by 
stinging them and storing them up as food for its young to 
illustrate the undeviating and unerring character of in- 
stincts. But the extensive studies of these entomologists 
led them to write : "The one preeminent, unmistakable and 
ever-present fact is variability. Variability in every par- 
ticular in shape of the nest and the manner of digging 
it, in the condition of the nest (whether closed or open) 
when left temporarily, in the method of stinging the prey, 
in the degree of malaxation, in the manner of carrying the 
victim, in the way of closing the nest, and last, and most im- 
portant of all, in the condition produced in the victims by 
stinging." 7 

No present-day authority so far as I know contends that 
instincts operate in a hard-and-fast manner comparable to 
the workings of any man-made machine. They are now 
universally recognized to be subject to the same general 

The Unity of the Organism 

principles of variation to which all organic phenomena are 
subject. Furthermore, under the searching investigation 
and criticism of numerous workers, notably H. S. Jennings 
and his followers, the tropism theory has been deprived, for 
most biologists, of its inorganically mechanistic character. 
The principles of "random movements," "avoiding reac- 
tions," "trial and error," and others, are thoroughly estab- 
lished and the recognition of them may be said to have so 
modified the doctrine of tropisms as to make it one of or- 
ganic mechanism rather than of inorganic mechanism as it 
virtually would be according to the thoroughgoing elemen- 
talistic conception of it. The "mechanistic conception of 
life," one may remark, has very much to commend it if only 
the machines conceived are recognized to be alive. My re- 
marks under this head * may be consulted by the reader 
who wishes to follow this point. 

What is meant by random movements is made clear by 
the following: "In the earthworm and the larvae of blow- 
flies which are negatively phototactic it has been shown by 
the writer that movements which bring the animal toward 
the light are checked or reversed and only those which hap- 
pen to direct the animal away from the light are followed 
up. Whatever immediate orienting tendency the light may 
have in these cases is relatively unimportant as compared 
with the element of selection of favorable movements in di- 
recting the animal away from the light." 

Here it will be noticed that the end, beneficial to the ani- 
mal, is reached through a combination of orienting reac- 
tions of the rigidly tropistic type, i.e., the type dependent 
on the movement of the animal directly toward or away 
from the source of light by the symmetrical plan of the 
body, and a sort of reaction in which the particular body- 
form and the direction of light rays are of only secondary 

* See "Machines, living," in the index of The Probable Infinity of Na- 
ture and Life. 

Organic Connection Between Physical and Psychical 253 

significance. But this latter type of activity, wholly di- 
vorced from a direct-orienting reaction, and even from a 
bilateral body symmetry, is of wide application among the 
lower animals. It was first brought clearly to the atten- 
tion of biologists by Jennings in his now well-known inves- 
tigations on Paramecium and other protozoans. These in- 
vestigations formed the bases of the "avoiding reaction" and 
the "trial and error" conceptions now generally recognized 
to be of much importance in the behavior of all animals, es- 
pecially of those in which a high measure of bodily activity 
occurs but in which there is little or no intelligence. Jen- 
nings' lucid account of his results in the chapter Be- 
havior of the Infusoria; Paramecium (Behavior of Lower 
Organisms) is strongly commended to the reader. 

The following paragraph must suffice for our reference 
to this work. After describing the behavior of Parame- 
cium, Jennings writes : '"This method of behaving is per- 
haps as effective a plan for meeting all sorts of conditions as 
could be devised for so simple a creature. On getting into 
difficulties the animal retraces its course for a distance, then 
tries going ahead in various directions, till it finds one in 
which there is no further obstacle to its progress. In this 
direction it continues. Through systematically testing the 
surroundings, by swinging the anterior end in a circle, and 
through performing the entire reaction repeatedly, the in- 
fusorian is bound in time to find any existing egress from 
the difficulties, even though it be but a narrow and tortuous 
passageway." And this complex and highly useful be- 
havior is performed by an organism which, so far as the 
best anatomical researches have been able to determine, is 
entirely devoid of a nervous system, and consists of a single 
cell ! 

But the "trial and error" scheme here exemplified is by 
no means confined to unicellular, non-nervous animals, nor 
to experimentally produced conditions. That it is opera- 

The Unity of the Organism 

tive in nature, and among animals with rather highly de- 
veloped nervous systems I shall illustrate by describing 
briefly a performance witnessed by me some years ago. 
This was the capture and engulfment of food by a nemertean 

These marine worms are of considerable size, some reach- 
ing a length of many inches, even a few feet, and ranging 
in thickness from less than an eighth of an inch to nearly an 
inch. Externally they give the impression of being very 
lowly in organization, the body being devoid of limbs or 
other appendages, and without segmentation. However, 
when they are examined internally a surprisingly high grade 
of organization is found, the muscular, digestive, blood and 
nervous systems being on a par, probably, with those of 
any invertebrates below the crustaceans and insects. The 
nervous system, particularly the brain, is relatively large, 
though not differentiated into diverse ganglionic masses and 
connecting strands to the extent found in jointed worms. 
The creatures are poorly equipped with external sense or- 
gans, there being no tentacles nor any certain olfactory or 
auditory organs. And eyes, when present, are so minute 
and simple as to be without power of sight in the ordinary 
sense; almost certainly they are mere light-perceiving or- 

The most distinctive anatomical feature of the nemerte- 
ans is a very long and thin though muscular and flexible hol- 
low tube situated at the anterior end of the animal, which 
is usually carried stowed away in a pouch within the body. 
While thus retracted the tube has some such relation to 
the rest of the animal that a glove-finger would have to the 

* Greatly to my regret I am unable to say what the species or even the 
genus was of either the nemertean or the annelid here referred to. The 
observation was made at the Shumagin Islands, Alaska, and under cir- 
cumstances that rendered it quite impossible to "look up" the species. 
And my knowledge of the taxonomy of these groups of worms is alto- 
gether too meager to enable me to identify genera even, offhand. 

Organic Connection Between Physical and Psychical 255 

glove were it completely inverted into the hand of the 
glove. This tube is used in the capture of prey, the ani- 
mals being carnivorous and highly voracious. The mode of 
employing the apparatus consists essentially in thrusting 
the tube out with almost the speed of lightning, the object 
being to bring the organ into contact with the prey at many 
points. The lash is not used as a lasso for catching or as 
a spear for piercing the prey, but for paralyzing it, prob- 
ably by a toxic secretion spread over the whole surface. 
The more effectually to accomplish this, the lash is shot out 
at a victim again and again. 

Now for the aspect of the whole operation of food-taking 
which specially concerns us. Being quite sightless and 
touchless in the usual sense, the lash must be used as an ex- 
ploring or finding as well as a paralyzing or killing organ ; 
and since its great length and limberness preclude it from 
being used as an ordinary tentacle is used, the finding op- 
eration is accomplished by repeated out-thrustings of the 
tube. In the instance witnessed the prey was an annelid 
worm, a creature well provided with locomotor organs, and 
a good crawler. On this account the victim-to-be was able, 
in the early stages of the onset, to move out of contact with 
the nemertean now and then. At such times the prey could 
be relocated only by darting out the lash at random, except 
as to general direction. So it resulted that many of the 
thrusts missed the mark; but they were instantly repeated 
with a little variation of direction, till the victim was lo- 
cated again. The whole performance reminded one of the 
game of blind-man's-buff, a game in which the seeker paws 
around in the general vicinity, as he believes, where the one 
sought was last touched. 

The effectiveness of the try, try again method was at- 
tested in this instance by the fact that the annelid was hit 
and the paralyzing dose administered times enough to put 
the annelid into so helpless a state that the nemertean was 

256 The Unity of the Organism 

finally able to get its mouth into contact with its prey. 
Then the victim, itself but little smaller than the nemertean, 
disappeared down the latter's "throat" with almost the 
rapidity with which the lash was retracted into and thrust 
out of its pouch. How much of this highly complex per- 
formance, so eminently useful to the nemertean, was purelv 
reflex, how much chemotactic, and how much instinctive { 
And who will assert positively that there was no trace of 
consciousness, even of intelligence, in it? 

An extremely interesting line of inquiry is suggested by 
cases of "trial and error" like this where at one extreme the 
"errors" are not much less numerous than the successes, and, 
at the other extreme, are cases in which the errors are re- 
duced almost to nil. A type case of this last would be the 
poise-and-spring of a cat after its prey. With little doubt 
a closely graded series could be made out running through 
from one extreme to the other. A cardinal interest in the 
inquiry would be as to the extent to which the simple reflex, 
tropistic reflex, instinct, and intelligence figure in the dif- 
ferent grades. Would it not turn out that the gradual 
diminution of error through the series would be, generally 
speaking, concomitant with the increase of intelligence? I 
suspect so. 

Tendency of Subrational Activities to Excessiveness 

The fourth and last certainty about reflex and instinc- 
tive activities to receive attention is their tendency to ex- 
cessiveness their way of going beyond what is necessary or 
even really safe for the welfare of the organism. Although 
from several points of view this is one of the most impor- 
tant aspects of the whole subject, it has received surpris- 
ingly little attention, especially by the modern school of ex- 
perimental zoology. 

Probably every one who has observed animals widely and 

Organic Connection Between Physical and Psychical 57 

thoughtfully has been impressed with the exuberance of their 
performances. That they are ever wont to overdo things, 
even operations which are when done in measured fashion 
absolutely essential to their existence, is matter of common 
knowledge. Holmes has some comments under this head 
which may fitly introduce our presentation. "With all their 
wonderful adaptiveness instincts are far from ideally per- 
fect. Much of Mark Twain's remarks on the futility and 
imbecility, the wasted effort and labor at cross purposes 
shown in the behavior of ants may easily be verified by any 

A common form taken by excessiveness of action is repe- 
tition. Very many, perhaps all, animals are notorious re- 
peaters. A few out of the many available instances will 
suffice to fix the phenomenon in mind. Some time ago a 
small whale (probably a half-grown Humpback, Megaptera 
versabilis) came near shore at La Jolla, California, and re- 
mained in the same small area for days. While there it 
went through a particular set of movements known to whal- 
ers as "breaching" scores of times, each set being exactly, 
so far as one could see from the shore, like every other set. 
The performance consisted, in this case, of a leap out of 
the water, which carried the body clear of the surface of 
the sea, the direction of emergence being probably thirty 
degrees from the perpendicular. During the ascent the ani- 
mal turned with a characteristic twist to the left and came 
down on its head and left side with a great splash. Once 
back in the ocean the creature reversed the course it was 
going when making the leap, returned to some distance from 
where it had emerged, reversed its course again, and re- 
peated the leap identically, to all appearances, even as to 
the spot of emergence and direction of travel. Why so 
many times the same performance in the same spot? That 
is the problem which concerns us here. Even though we 
conceive it to be somehow adaptive connected in some in- 

258 The Unity of the Organism 

direct way possibly with feeding or reproduction or migra- 
tion or some other vital function the question still remains, 
why so much of it? And to this no probable or even ra- 
tional answer is forthcoming from the standpoint of adapta- 
tion and utility, taking these terms in their usual meaning. 

Here is another case from the mammalia, the possible 
adaptive significance of which is still more remote, if any- 
thing, than that of the behavior of the whale. Many indi- 
vidual mice of the genus Peromyscus being used by Doctor 
Sumner and Mr. Collins in their researches on heredity and 
environmental influence at the Scripps Institution take to 
throwing back summersaults in their cages. The more com- 
mon performance consists in a run along the floor of the 
wooden cage and up its side to near the top, then a quick, 
strong jump backward clear across the cage, the feet being 
uppermost during the first part of the leap but coming to 
rights again by the time the landing is made. Here again 
the question of why the mice do this seemingly useless thing 
is not so interesting for the present discussion as that of 
why they do it so much. 

The high flight of some species of birds, the great eleva- 
tions being reached by long, regular upward spirals, would 
appear to come under the head of non-adaptive, superfluous 
action. The sand-hill crane, Grus mexicana, may be taken 
as an instance of a bird given to this habit. Surely such 
flights by this species can have nothing to do with food- 
getting, since in the excursions the bird is going directly 
away from, instead of into, the region where its food 
abounds. It eats snakes, frogs and other creeping animals, 
and various seeds and roots. Nor is there any evidence that 
the flights are concerned with the mating function, nor yet 
with migration, though one might possibly imagine that 
while on the excursions the birds learn, after a fashion, the 
topography of the surrounding regions. 

The high-diving and booming of the night-hawk, Cordeiles 

Organic Connection Between Physical and Psychical 259 

liry'ununiis, repeated time after time in the early evening and 
occasionally in midday when an approaching storm cools 
the air, would seem to be another performance of the non- 
adaptive sort. The suggestion that this is a courtship af- 
fair can hardly stand, in view of the fact that at least as 
often as otherwise the birds which do it are entirely alone. 
Xor can one see how so extensive and swift a dive, with so 
much noise, can be advantageous for the capture of flying 

And reflect on the quantity of movement of many ani- 
mals. Can any one believe that mammals and lizards run, 
birds and insects fly, and fishes swim just exactly so much as 
and no more than, they must in order to survive? Would 
it be contended that the Golden Plover, to take a well known 
case of extensive migration, would certainly succumb in the 
struggle for existence on anything less than a journey from 
the high latitudes of the northern hemisphere well into the 
southern hemisphere and back, each year? There is a vast 
difference betw r een a necessity for migration to some ex- 
tent and a necessity for migration of a particular quantity. 
One of the great weaknesses of the natural selection theory 
has been, I am very sure, its slight regard for quantity ; 
quantity of need, quantity of performance, quantity of 

These examples serve to illustrate the fact that among 
the higher animals at least, much muscular activity occurs 
which is not at all, or only partly, adaptive. But by far the 
more common occurrence of excessive activity is in connec- 
tion with behavior which is more or less obviously adap- 
tive. "A good thing carried to excess," in the familiar 
phrase, expresses w r ell what is in mind here. 

This excessiveness of adaptive activity is naturally more 
easily recognized in animals which are most easily observed 
and most active generally. Thus it is from birds and in- 
sects that examples can be most readily drawn. 

260 The Unity of the Organism 

Let the current view be accepted that the song of pas- 
serine birds is associated adaptively with the mating func- 
tion. Even so, no one who has given careful attention to 
the matter can have failed to recognize that with many 
species much more singing is done than actual pairing and 
breeding call for. I have kept almost daily notes for sev- 
eral years on the singing of the Western Meadow Lark, 
Sternella magna neglect a, in the vicinity of La Jolla. The 
birds are resident the whole year through, and as they come 
familiarly around my home and laboratory, the observations 
can be quite full. Although the breeding time is restricted 
to late February, March, April, and sometimes May, there 
is not a month in the year when songs may not be heard, 
most of the time in full volume. Significantly, I believe, the 
song is at its ebb during some weeks just before the nesting 
period begins. Nor does the singing of the males seem to 
be connected in any close way with mating. The birds do 
not pair off closely and permanently, even for the breeding 
season. Most of the singing, which occurs chiefly in the 
morning and early forenoon and again toward evening, is 
done while the singer is, more commonly than otherwise, 
quite alone on some telephone pole or wire. And the mode 
of singing does not change at all when mating begins. An- 
other interesting fact about the singing of this species is 
the considerable range of temperature and light conditions 
over which the song is invariable, so far as these factors are 
concerned. The song may be as full and frequent on 
cloudy, misty mornings as on sunny ones ; and over a con- 
siderable range of temperature the song is quite independent 
of the particular degree marked by the thermometer. 

While the song habits of this bird are undoubtedly some- 
what exceptional in their looseness of correlation with mat- 
ing and with environmental conditions, certain it is that 
much this sort of thing is observable with several resident 
species which I have observed. The house finch, Carpodacus 

Organic Connection Between Physical and Psychical 261 

Mexicanus, and the California towhee, Pipilo fusc-us, may be 
specially mentioned in this connection. The fact that do- 
mesticated song birds, like the canary, may be brought to 
sing almost perpetually is only an extreme manifestation 
of tendency among song birds to sing in excess of any strict 
utility of song. 

Think of the monotonous repetition in the croaking of 
frogs, the chirping of crickets, the stridulations of cicadas, 
and so on ! I have counted more than five hundred con- 
secutive chirps of a cricket in about half an hour, with 
only a little variation as to notes or intervals. And this is 
surely a very moderate example of what actually occurs- 
as any one can easily convince himself by listening and count- 
ing almost any still night, almost anywhere where crickets 
live. Probably the chirping of crickets is employed in mat- 
ing. Very well. But are the thousands of chirps uttered 
by a given individual each night for many nights, the small- 
est number upon which the species can survive? Even ask- 
ing of the question reveals the monstrosity of a theory that 
would necessitate an affirmative answer to it as strict ad- 
herence to the natural selectionist meaning of utility un- 
doubtedly would. 

In place of bringing forward additional instances, which 
could easily be done, to show that vocal sounds and bodily 
performances of various sorts more or less obviously con- 
nected with mating among higher animals are produced in 
excess of what the strict application of the rule of physio- 
logical economy would dictate, I shall do no more than util- 
ize the conclusions of two investigators who seem specially 
qualified to speak on the subject, and assume that these con- 
clusions would receive the sanction of all zoologists who have 
given serious attention to the matter and have formed their 
judgments unbiased in favor of any explanatory theory. 

The first of these investigators is W. H. Hudson, who 
represents a period a little antecedent to the present spe- 

262 The Unity of the Organism 

cially critical experimental era. I quote from his well- 
known The Naturalist in La Plata, published in 1892: "I 
wish now to put this question: What relation that we can 
see or imagine to the passion of love and the business of 
courtship have these dancing and vocal performances in 
nine cases out of ten? In such cases, for instance, as that 
of the scissor-tail tyrant-bird, and its pyrotechnic evening 
displays, when a number of couples leave their nests, con- 
taining eggs and young, to join in a wild aerial dance; the 
mad exhibitions of ypecahas and ibises, and the jacanas' 
beautiful display of grouped wings ; the triplet dances of 
the spur-winged lapwing, to perform which two birds already 
mated are compelled to call in a third to complete the set; 
the harmonious duets of the oven-birds, and the duets and 
choruses of nearly all the wood-hewers, and the wing-slap- 
ping aerial displays of the whistling widgeons ; will it be 
seriously contended that the female of this species makes 
choice of the male able to administer the most vigorous and 
artistic slaps? . . . There are many species in which the 
male, singly or with others, practises antics or sings during 
the love-season before the female ; and when all such cases, 
or rather those which are most striking and bizarre, are 
brought together, and when it is gratuitously asserted that 
the females do choose the males that show off in the best 
manner or that sing best, a case for sexual selection seems 
to be made out. How unfair the argument is, based on these 
carefully selected cases gathered from all regions of the 
globe, and often not properly reported, is seen when we turn 
from the book to Nature, and closely consider the habits and 
actions of all the species inhabiting any one district. We 
see then that such cases as those described and made so much 
of in the 'Descent of Man,' and cases like those mentioned 
in this chapter, are not essentially different in character, 
but are manifestations of one instinct, which appears to be 
almost universal among the higher animals. The explana- 

Organic Connection Between Physical and Psychical 263 

tion I have to offer lies very much on the surface. . . . We 
see that the inferior animals, when the conditions of life are 
favorable, are subject to periodical fits of gladness, affecting 
them powerfully, and standing out in vivid contrast to their 
ordinary temper. And we know what this feeling is this 
periodic intense elation which even civilized man occasionally 
experiences when in perfect health, more especially when 
young. There are moments when he is mad with joy, when 
he cannot keep still, when his impulse is to sing and shout 
aloud and laugh at nothing, to run and leap and exert him- 
self in some extravagant way." 

The reader is asked to note what Hudson says about pick- 
ing out such evidence as will help the case for sexual selec- 
tion, and saying nothing about evidence which will not help 
it. Beyond question the dogma of natural selection, espe- 
cially the Weismannian perversion of it, has flourished 
largely on this sort of thing. Nor has natural selection alone 
among biological theories had the benefit of assorted evi- 
dence. Indeed the whole elementalistic mode of interpret- 
ing living nature may be characterized as one whose doc- 
trines depend largely upon "special privilege," to adopt a 
phrase lately much used in the economic world, as to evi- 
dence for their support. 

The other investigator upon whom we draw is Prof. Julian 
S. Huxley, whose work is that of a field zoologist imbued 
with the exacting spirit of the present day. Huxley's stud- 
ies are devoted to the mating habits of birds, so there can 
be no question that the activities he describes are intimately 
connected with reproduction. Of the numerous species 
dealt with in the paper now before us, we notice first the 
Great Crested Grebe. It is highly significant that in this 
species mating takes place before the so-called courtship 
performances begin, so this latter process can not be es- 
sential to securing a mate. The female is "courted" after 
she is got possession of. The courtship activities begin soon 

The Unity of the Organism 

after pairing, two entirely different sets of ceremonies be- 
ing involved in the activities. One of these Huxley calls 
ceremonies of mutual display, the other, ceremonies of coi- 
tion. The highly elaborate mutual display performances 
are fully described but can not be reproduced here. They 
consist in a variety of body attitudes, head and wing and 
feather movements, swimmings and divings, and call-notes, 
the whole lasting some minutes. Concerning this prelim- 
inary operation, Huxley writes : 

'The most noticeable thing about all these ceremonies is 
that they are 'self-exhausting'- -they do not lead on to any- 
thing further. Looked at from the physiological point of 
view, they seem to me to be nothing but 'expressions of emo- 
tion' : the birds act thus because they are impelled to do 
so, because they enjoy it. Looked at, on the other hand, 
from the evolutionary point of view, they seem to have been 
developed as a bond to keep the pair together." 12 

Following these preliminaries, the ceremonies of coition 
take place, these being less striking, though characteristic. 

Speaking of his studies on the mating habits of some of 
the warblers, and referring to differences of interpretation 
between himself and W. P. Pycraft, another observer in the 
same field, Huxley writes : "In this, Mr. Pycraft and myself 
are, I think, agreed ; to both of us the 'display' of the male 
Warbler is nothing but a direct expression of sexual excite- 
ment, scarcely, if at all, modified by Darwinian Sexual Se- 
lection nothing but the way in which nervous disturbance 
caused by sexual excitement happens to liberate itself. Gen- 
eral nervous discharge will cause general muscular contrac- 
tion ; and something approaching this is here seen rapid 
hopping, extension and fluttering of the wings, spreading 
of the tail, bristling up of the feathers on head and 
throat, and utterance of a series of quick sounds. This ex- 
presses a condition of readiness to pair, and doubtless to 
the female comes to be a symbol of the act of pairing. 

Organic Connection Between Physical and Psychical 265 

Hence, as far as the female is concerned, the act of pairing 
has come to depend upon this stimulus (acting of course on 
a suitable internal physiological state). This is no more 
strange in the bird than it is that in ourselves thoughts and 
emotions of love well up at the sight of some tangible ob- 
ject connected with the beloved." 13 

But it is in the sex function itself that the tendency to 
overdo manifests itself with greatest force. In fact, the fa- 
miliar and ominous expression "sexual excesses" as applied 
to the human animal, indicates very truthfully w r hat is be- 
fore us. The whole phenomenon of competing and fighting 
among the males of all higher animals for possession of the 
females, with its momentous consequences in dozens of ways, 
may truly be said to rest back on the excessiveness of the 
sex impulse and instinct. Since as a general rule the males 
and females of animal species are approximately equal in 
numbers, pairing off two by two, after the manner of the 
population of Noah's ark, might occasion but little and mild 
competition could each male and each female be satisfied 
with one mate, in accordance with the allotment which the 
numerical equality would make. And the pertinent question 
may be raised in passing, would not such a mode of pairing 
secure the perpetuation of the species quite as well as, pos- 
sibly better than, the method which is so largely in vogue? 

Highly suggestive seems to me in this connection, observa- 
tions I have recently been able to make on the mating hab- 
its of one of the California "surf perches" (Cymatogaster 
aggregatus). This is one of the numerous viviparous bony 
fishes peculiar to our coast. The species under attention 
lives quite normally, as far as one can see, in the aquaria of 
the Scripps Institution ; so what may be assumed to be its 
typical habits can be observed continuously. 

Strict monogamy appears to prevail in the species. At 
least this is true with the specimens three males and four 
females under observation, and so far as a particular breed- 

266 The Unity of the Organism 

ing period is concerned. Each male begins his attentions 
while his fiancee, so to speak, is heavily gravid from the pre- 
vious mating (when and how accomplished we unfortunately 
know nothing about beyond the fact that it must have been 
before the individuals under observation were brought to 
the aquarium from the sea, about six weeks before the mat- 
ing began. ) 

In the case of one pair, the amours of the male continued 
more than two weeks, the first few days of which were 
before the family of young began to be born, the period of 
parturition extending over three days. Although there 
was no indication on the part of the other males of inten- 
tions or even desires toward the spouse (as she may now 
be called) of this male, he was quite pugnacious, directing his 
seemingly unnecessary operations against the other females 
as well as against the other males. It should be said, how- 
ever, that his antipathies were considerably greater against 
the males than against the other females. The other two 
males took partners after much the same fashion ; but since 
both of these were somewhat smaller, and fully acknowl- 
edged the over-lordship of the one singled out in our account, 
their performances were less clear cut. 

Specially noteworthy is the character of the amours of 
the male, which alone or almost alone, seems to take an in- 
terest in the performance. No contact, or at least only the 
slightest, of the male with the female was seen though the 
fish were under observation much of the time. A peculiar 
downward darting of the male first on one side then on the 
other of the female, close to her but not quite touching her, 
was one of the favorite manoeuvers. But various rapid 
circlings about, up and down, head-on and tail-on, over and 
under, and in nearly all possible ways, may be witnessed. 

The full meaning of this monogamic (temporarily so, at 
least), largely non-tactual type of mating we do not know 
partly because we have not yet all the facts ; but I suspect 

Organic Connection Between Physical and Psychical 267 

it to be important. But this much is clear as to its bear- 
ing upon the point uppermost in this discussion: There is 
an excessiveness of activity in a variety of ways, particularly 
in the driving of other females, the presence of which in the 
vicinity of the mate is merely incidental and utterly harm- 

Obviously it is the demand, instinctive or organic or both, 
for more sexual gratification than the natural numerical 
scheme of the two sexes provides, and the actual necessities 
of race perpetuation demand, which is largely responsible 
for the contests to secure mates, so characteristic of all 
higher animals. The bull fur seal must have forty or fifty 
mates, instead of the one which the numerical equality of 
the two sexes would naturally give him ; hence the fierce com- 
bats among the males, with the result that a great majority 
of the whole male population at any one time is forced to 
remain outside the "harems" during the mating season. And 
some such eliminative process must occur in all species where 
the sexes are about equal in numbers, and where promiscuity 
in pairing is practised. 

Nor are the injuries and disasters which may result from 
the driving power of the sex-impulse restricted to compet- 
ing individuals of the same sex. The mates sought after not 
infrequently suffer seriously from the excesses of the seek- 
ing males, the females being usually more passive and hence 
the more liable to injury in this way. Thus, J. S. Huxley 
has lately told of the exhaustion and actual death of the 
female mallard duck from being repeatedly "tread" by the 
males, the same and different individual males participating 
in the strangely destructive performance. 

Finally, the individual itself is not safe from self-injury 
through its own sex impulses. Some of the forms which this 
sort of thing may take in the human species are too famil- 
iar, too disastrous and too repugnant to need illustration 
in proof of their reality. That they occur also more or less 

The Unity of the Organism 

among animals is well known to all who have had consid- 
erable experience with domestic animals. 

Excessive activity in connection with the alimentary func- 
tion must now be glanced at. That there is no nice quanti- 
tative balance between the food necessities of the animal and 
the food gathering instincts and impulses and efforts on the 
basis of the principle of natural economy and parsimony, 
is shown conclusively it would seem by many animals which 
have the storing habit. The honey bee is an example of 
this among insects. Given a sufficient supply of flowers to 
work on, in the wild state these bees seem always to store 
away more food material than they consume. 

The extent of their honey-making is limited rather by 
the raw material available and by their own restricted phys- 
ical powers than by their nutritional needs. This is the im- 
pression I have from my observations on wild and tame bees 
and I find it to coincide with that of other naturalists whose 
opportunity for observing wild bees has been much greater 
than mine. For example my esteemed naturalist friend, Mr. 
Frank Stephens of San Diego, California, reminds me that 
the view is confirmed by the fact that in "bee trees" a por- 
tion of the comb containing honey is not infrequently black 
and shows signs of being old. 

Darwin made quite a point, it may be recalled, of the 
economy in some aspects of the bee's work. "The comb of 
the hive-bee," he says, "as far as we can see, is absolutely 
perfect in economising labour and wax." (Cell-Making In- 
stinct of the Hive-Bee, in The Origin of Species.) 14 But a 
thoroughly economic adjustment between different parts of 
a given complicated operation, and economy of the opera- 
tion as a whole, are very different. 

As an instance of excessive repetition in the food-getting 
activities among the insects, the following from Fabre may 
be taken as fairly typical. A solitary wasp of the genus 
Sphex captures and slays a locust, but instead of using it 

Organic Connection Between Physical and Psychical 269 

at once for food, or of taking it directly into her home, she 
sometimes leaves it on the road, and runs to her home, even 
though this is threatened by no danger. Then after a time 
she returns to the game. This going-and-coming may be 
performed repeatedly before the carcass is finally taken into 
the dwelling. If by chance the game is removed during the 
absence of the wasp, the wasp returns to the spot where her 
load was left, but, not finding it, she, nevertheless, keeps up 
the going-and-coming for some time. The first back-and- 
forth journey from game to dwelling is explicable, Fabre 
shows. "But what is the use of the other visits, repeated 
so speedily one after another?" Fabre inquires. 1 Something 
like this almost every one must have seen, who has watched in- 
sects at all. 

I am quite certain that the acorn storing habit of the 
California woodpecker, Melanerpes formicivorus bairdi, is 
quite beyond any use the bird makes of the acorns. In the 
first place, despite much discussion of the question whether 
the acorns are used at all, and if so how, the case is by no 
means clear. But the point I particularly wish to make is 
that whatever use, if any, the birds make of the acorns, 
whether as food directlv or as culture media for worms or 


insects, these in turn to be eaten by the birds, they store up 
many more than they utilize. This seems to me highly prob- 
able from the fact, which I have ascertained by numerous 
examinations at different places and times, that many holes 
contain dried up and wasted acorns which show no signs 
of having been picked at or otherwise moved after they were 
inserted into the holes. Furthermore, the great extent of 
the hole-drilling and filling in itself seems to exceed the 
bounds of necessity, especially in view of the certainty that 
the bird's chief food supply is from quite another source. A 
pine log fifty feet long and one hundred thirty-six inches in 
girth at the middle, which I found in the San Jacinto Moun- 
tains, contained on a fairly careful estimate 31,800 holes, 

270 The Unity of the Organism 

many of them containing acorns. 

But even were it certain that the acorns are utilized in any 
manner and to some extent in connection with the feeding 
function, there are still other evidences than that just ad- 
duced of the imperfect and excessive operation of the acorn- 
storing instinct. As is well known, the bird sometimes ex- 
tends its drilling operations to wooden buildings to the ex- 
tent of making itself a great nuisance. I have seen a case 
where the birds had pierced the rustic of an uninhabited 
house, so that when the acorns were inserted, instead of 
filling the puncture as they would fill holes in a tree, they 
would drop down into the space between the rustic and the 
inner wall. Apparently the failure to stop the hole, and 
failure also to perceive why, or to recognize that the hole 
could not thus be stopped, "fooled" the birds into putting 
one acorn after another into the same hole, endlessly almost, 
judging by the great quantity of nuts piled up at the bot- 
tom of the space. 

While the storing habit of the California woodpecker is 
undoubtedly exceptional as to extent, it is by no means 
wholly unique. At least one species of blue- jay (Cyanocitta 
cristata) has much the same habit, in the opinion of most 
ornithologists who have studied the habits of the bird. An 
experienced naturalist, E. H. Forbush, has recently said 
concerning Mark Twain's "Baker's Blue Jay Yarn," in A 
Tramp Abroad, "All of this is not merely amusing; it is good 
ornithology in so far as it reports the way a Jay acts." 
This story, it may be said for the benefit of any reader so 
unfortunate as not to know it, turns upon the performance 
of a jay similar to that narrated above about the California 
woodpecker, the acorns, and the old house. 

The habit of the shrikes (genus Lanius) of impaling their 
victims and leaving them, almost certainly operates more or 
less independently of, and often in excess of, the food re- 
quirements of the birds. "My observations," says Forbush, 

Organic Connection Between Physical and Psychical 271 

"have led me to believe that it rarely returns to eat what it 
has thus cached, unless driven to do so by hunger resulting 
from adverse fortunes of the chase." 17 

Nor is there much if any question that something of the 
same sort occurs among mammals which have the food stor- 
ing habit. E. T. Seton quotes the following from Dr. John 
Wright concerning the big eastern chipmunk (Tamias stria- 
tus griseus) : "It is a most provident little creature, con- 
tinuing to add to its winter store, if food is abundant, until 
driven in by the severity of the frost. Indeed, it seems not 
to know when it has enough, if we may judge by the surplus 
left in the spring, being sometimes a peck of corn or nuts 
for a single squirrel." There are many other statements 
by the best authorities, especially concerning numerous spe- 
cies of mice, which strongly suggest a like superabundance 
of storing activities. But for the rest I will mention a case 
that has come to my own notice. 

I am indebted to Mr. Frank Stephens for information 
about and the opportunity to witness to some extent for 
myself the operations of the storing instinct and feeding 
habits of the Antelope Ground-Squirrel (Ammosperinophi- 
lus leucurus). This chipmunk-like little squirrel proves to 
be so readily domesticable that it becomes almost as familiar 
a household member, at least for Mr. Stephens' household, 
as a domestic cat. Although an account of the habits of the 
single individual in Mr. Stephens' possession can not yet be 
told fully by a long ways, a few points of much interest 
for the present discussion are positive enough. 

In the first place the genuinely instinctive character of 
the storing habit is established by the fact that although the 
specimen under observation was taken soon after birth, and 
has lived all its life in complete isolation from parents and 
all its kind and has been furnished artificially with an 
abundance of food, its storing operations are carried on 
constantly and almost as perfectly, so far as one can judge. 

The Unity of the Organism 

as though it were living in the natural state. This fact in 
itself is evidence that the instinct is not determined solely 
by immediate needs of the individual. But much more 
convincing evidence furnished by this case to this effect is in 
the particular way the instinct works. For example, this 
species possesses cheek pouches for carrying food as do so 
many rodents which have the storing habit. When nuts, 
grain, etc., are presented to the animal she very rarely eats 
them immediately even though manifestly hungry, but carries 
them away to some distance, one at a time ; going back and 
forth and placing the articles in her two pouches till these 
are quite full. And these little pre-storage journeys, as 
they may be called, are often definite in character. At any 
given time they end at nearly the same spot, and the animal 
takes nearly the same position while the article is being pre- 
pared for and inserted into the pouches. This is clearly 
the typical procedure in filling the pouches, though it is 
varied considerably from time to time. 

As to what follows the pouch-filling there is considerable 
variation normally so it appears. In case the animal is 
hungry she may quietly extract the nuts from the pouches 
and eat them. Or she may run about for some time with 
her cheeks bulging full. Or she may take her load off some- 
where and lay it away either in some cache previously es- 
tablished or in a new one. The cache may be in a bed of sand 
if this is at hand ; or it may be in or under some old garment 
or piece of cloth or paper which the surroundings may 

An especially interesting fact noticed by Mr. Stephens 
is the tendency shown on the part of this squirrel to carry 
the articles to as distant a place from where it gets them 
as can well be reached. 

On the whole there is no doubt that we have here a var- 
iedly illustrative example of activity over and above need 
in the operation of an instinct. 

Organic Connection Between Physical and Psychical 273 

This bare touch, so far as instances are concerned, of 
overactivity in connection with reflexes, and especially with 
instincts which are on the whole useful, leads naturally to 
the great field of animal play. Space limitations prohibit 
us from taking more than a bird's eye view of this field. 
Fortunately, however, even such a view can be quite effective 
for our purpose because of the well-known work of Karl 
Groos, The Play of Animals. Our sole purpose here, as 
in the rest of this discussion, is to answer the question 
whether animals do or do not carry their activities which on 
the whole are fundamental to their existence beyond what 
is necessary for their own individual requirements. With 
Groos's explanatory theory of play we are concerned only 
so far as it involves the question of fact upon which our 
present interest centers. That most if not all animal activ- 
ity which can rightfully be called play, and which is not 
intelligent, is instinctive, we believe Groos has conclusively 
shown. The explanation adopted by Spencer and others 
that play is the useless imitation by young animals of useful 
activities performed by their seniors, the imitative acts be- 
ing useless because merely the overflow of "surplus energy," 
is certainly inadequate, as Groos has insisted. That animals 
constantly go through performances playfully which they 
have had no chance to see or to have otherwise impressed 
upon them from without, is as certain as that they constantly 
perform useful acts in this way. 

It consequently results that a source of energy for play, 
that is, for actions which are not immediately essential to 
the existence of the organism, must be an endowment of the 
organism no less certainly than that a source of energy must 
exist for actions which are essential to its existence. So 
Groos's statement : "A condition of surplus energy still ap- 
pears as the conditio sine qua non that permits the force 
of the instincts to be so augmented that finally, when a real 
occasion for their use is wanting, they form their own mo- 

The Unity of the Organism 

tive, and so permit indulgence in merely sportive acts," 
becomes a statement of fact if by "surplus energy" we 
understand energy available for, and upon occasion used for, 
acts which are not indispensable to the existence of the in- 

The quantity and generality of play performed by animals 
may be taken as one important measure of the extent of the 
energy possessed over and above what is essential for their 
normal individual existences, and this without reference to 
whether or not the play may be useful as a preparation for 
future essential activities, or for recreation only. The fact 
can hardly be too much insisted upon that ulterior useful- 
ness of the organism's acts, whether to the species generally, 
to offspring, or to the individual's own future, cannot pos- 
sibly be a sufficient explanation of the energy immediately re- 
quired for the act itself. Even though an animal does noth- 
ing whatever except by reason of its hereditary endowments, 
or in the interest of its offspring; and though the real pur- 
pose of much that it does looks to its own future, it must 
nevertheless continue to eat, digest and assimilate, and 
breathe. The subdivision of biology which has come to be 
known as physiology has for its distinctive task exactly 
that of studying the present activities of the organism. With 
the organism's past, whether individual or racial, and with 
its future, whether individual or racial, physiology can be 
concerned only indirectly. 

Summary of Organismal Character of All Subrational 

Psychic Life 

Having now examined broadly though far from exhaus- 
tively the psychic life of the animal in each of its most ob- 
vious phases, the highest rational phase, the emotional phase, 
the instinctive phase, and the reflex phase (in which tropisms 
are included) for purely descriptive and classificatory pur- 

Organic Connection Between Physical and Psychical 275 

poses, let us briefly summarize what we have learned. 

In each phase we have found the organism, living, whole 
and normal, indispensable to a comprehension of the phe- 
nomena examined. Or, expressed in a different way, we have 
found it possible in each phase to reach only a very imperfect 
understanding of the phenomena by referring them to the 
elements which can be discovered in them. For example, the 
theory of association of ideas is inadequate to explain ra- 
tional life, in such manifestations as apperception and mental 
initiative and creativeness. 

In the emotional phase, in such emotions as fear, rage and 
sex passion, not only does cursory observation recognize 
the involvement of a large part of the organism, but physi- 
ological investigation is able greatly to extend our recogni- 
tion of this involvement by showing how the nervous system 
in its cerebro-spinal and its autonomic divisions, the cir- 
culatory, the alimentary, and the internal-secretory sys- 
tems, are essentially and reciprocally involved. 

As to the organismal character of psychic life in the 
phase of instinct, it suffices to recall that one of the most 
widely accepted criteria for differentiating instinctive from 
reflex activities is that the former involve the organism as a 
unity, a whole, while reflexes, according to this criterion, 
pertain only to limited portions of the organism. "An 
instinct is a more or less complicated activity manifested by 
an organism which is acting, first, as a whole rather than as 
a part." 20 To this statement of the matter may be added 
that when the instinctive act is in the interest of the indi- 
vidual performing it, the act is not only by but for the in- 
dividual as a whole. 

As to the reflex phase (if that is to be reckoned as psy- 
chic) the organismal nature of tropisms has had so large 
a place in our discussion that surely no more need be done in 
this summary than to remind the reader of our discussion of 
tropisms. And even reflexes of a simpler fom than the 

276 The Unity of the Organism 

tropisms indeed the abstract conception of the "simple 
reflex," though not, perhaps, involving the conception of the 
organism as a whole, yet is not comprehensible on elemen- 
talistic principles, as our examination of Sherrington's in- 
vestigations revealed. And such phenomena as those of the 
spreading and compounding of reflexes are quite incompre- 
hensible except on the organismal principle, even though the 
whole organism may not be involved, observably at least, in 
particular reflex acts. 

Specificity of Subrational Psychic Life 

The concluding section of this descriptive chapter on 
psychic integratedness must be devoted to the specificity, 
not to say individuality, of animal behavior in all its phases. 
The vast body of trustworthy detailed knowledge now in our 
possession justifies, I am quite sure, the following general- 
ized statement under this head: It is exactly on the psychic 
side of animal life, psychic being taken in the broadest sense, 
that animals are most differentiated from one another, both 
as to individuals and as to species. 

Taxonomic zoology is based almost entirely on structural 
attributes of animals. This results from reasons that are 
obvious, speaking generally, and constitutes a justification 
of the fact from a practical standpoint. Nevertheless the 
purely practical advantages of the classificatory systems as 
they have been built up have been, and are, gained at the 
expense of several rather serious disadvantages. One of 
these is, as advance of knowledge leads us to realize, that our 
well-nigh exclusive attention to structural differences and 
likenesses has tended strongly to divert attention from func- 
tional differences and likenesses. It is of fundamental impor- 
tance for a truly comprehensive science of organic beings, 
that is, for a philosophical biology, to regard our synoptic 
classifications not as a final result of knowledge-getting, but 

Organic Connection Between Physical and Psychical 277 

rather as a life-sized sketch, as one may say, of the whole 
living world, to facilitate the gigantic task of completing 
the picture through the cooperation of numberless artists, 
the completion to be accomplished by filling in the sketch 
with the entire round of attributes, structural and func- 
tional, presented by the natural lives of organisms. I have 
dwelt somewhat at length on this matter elsewhere,* and can 
refer to it here only as a background for what I wish to say 
about psychical specificity. 

Two extracts must suffice. "No biological phenomenon 
is adequately interpreted or dealt with experimentally, until 
it has been considered with reference to the place that the 
organism to which it pertains holds in the system of classi- 
fication." And further: "What I affirm is that the inductive 
evidence has now gone so far toward proving every sharply 
differentiated species to contain some differentia in all the 
main provinces of their structure and function, that to as- 
sume the absence of such differentia in any given case is 
unwarranted." 21 

I want to utilize these earlier general statements about 
organic specificity, as a basis on which to rest a generaliza- 
tion concerning the specificity of psychic attributes. So 
enormous is the observational data available for illustration 
here, that in lieu of presenting any of them I am going to 
state in a wholly dogmatic fashion the generalization toward 
which we are certainly being led by modern crucial researches 
on animal behavior. Let us imagine ourselves possessed of an 
approximately exhaustive descriptive knowledge of the be- 
havior of the whole animal world, this knowledge being, how- 
ever, unaccompanied by any knowledge whatever of the cor- 
poreal nature of the animals. This behavior-knowledge 
would fall naturally into categories larger and smaller, the 

* The Place of Description, Definition and Classification in Philoso- 
phical Biology, in The Higher Usefulness of Science (Boston, 1918) ; 
also The Scientific Monthly, November, 1916. 

278 Tlie Unity of the Organism 

smaller ones being for the most part subdivisions under the 
larger. Then let us imagine this system of behavior-knowl- 
edge compared exhaustively with a later-acquired, equally 
exhaustive knowledge of the corporeal nature of all animals. 
The two systems would be found to match each other very 
nearly as closely as though the two had been worked out to- 
gether, much as they are being actually elaborated by struc- 
tural and functional zoology. In other words, the species, 
genera, orders, and so on, of animals are differentiated from 
one another and coordinated with one another by their "be- 
havior," that is, by their whole round of psychical and reflex 
attributes, much as they are by their corporeal attributes. 

The inductive evidence for such a generalization is being 
produced at the present time by three quite definitely marked- 
off kinds of research. These may be designated as ( 1 ) quali- 
tative field researches, (2) laboratory experimental re- 
searches, and (3) quantitative field researches, the definitely 
quantitative method being statistical. 

The first-mentioned class of investigations is typified by 
the earlier field zoologists, whose aim was to learn, as ex- 
haustively as possible in a purely qualitative way, the habits 
of animals in nature. Workers of this class are the typical 
zoological naturalists of the history of animal biology. 
Aristotle, Conrad Gesner, John Ray, Charles Linne, P. S. 
Pallas, Gilbert White, J. J. Audubon, J. H. Fabre, A. R. 
Wallace and A. Forel may be named as conspicuous examples 
of pre-modern members of this class ; and Charles Darwin 
stands out sharply as a representative of it, but as a transi- 
tion to the modern period, the transitional character of 
Darwin being seen not only in the doctrines he proposed but 
as well in his intimate combination of the experimental 
method with the older method of observation. 

By the modern period of research in field zoology I mean 
the period during which, while the natural history point of 
view and attitude are retained, the critical rigor of modern 

Organic Connection Between Physical and Psychical 279 

science generally is practiced, and experimentation in one 
form or another is employed as a supplementary agency 
wherever and whenever possible. One of the best examples 
of this type of zoological research and writing is Ants, by 
W. M. Wheeler. But a considerable portion, and fortunate- 
ly an increasing portion, of experimental research in animal 
behavior is being done quite in the spirit of field zoology. 
The work of R. M. Yerkes deserves mention as perhaps the 
most definitely purposed and executed combination of the 
field and experimental methods for investigating the behavior 
of mammals and birds, that has yet been made. But much 
of the research recently named animal ecology tends strong- 
ly toward rigor in field investigation. This kind of study is 
specially adapted to bring out the specific nature of behav- 
ior, since the group of organisms, species, etc., as a whole, 
occupies a central place in the student's interest, so that if 
behavior is attended to at all its differential features are 
likely to receive attention along with the differential struc- 
tural features. 

(2) Laboratory experimentation on animal behavior has, 
as previously indicated, been prosecuted more intensely and 
widely in the modern period than either of the other classes 
of investigation. In fact, it may be said to be distinctive of 
the period, and to have set the standard as regards rigor for 
the other types of investigation. From its very nature, how- 
ever, it is not calculated to bring the specificity of behavior 
to a central place in the student's interest. Singling out 
as it does one or a few attributes at a time as they are ex- 
hibited by one or a few individuals of one or a few species, 
breadth and penetration of comparison are liable to be sacri- 
ficed. This kind of research tends to be extremely particu- 
laristic in every way. Nevertheless, painstaking and judi- 
cious workers, like Englemann, Forel, Binet, Wasmann, 
Romanes, C. Lloyd Morgan, Verworn, Jennings, Loeb, 
Holmes, and Parker, generally state what species their in- 

280 The Unity of the Organism 

vestigations have dealt with, so a reader interested in the 
question of specificity can usually detect evidences of dif- 
ferences in the behavior of different species, even though the 
investigator himself was obviously little interested in the 
subject, and so took no pains to bring out such evidence. 

Indeed, the fact that species-differences in behavior can so 
frequently be recognized in descriptions even though the 
writer's general attitude may, if anything, militate against 
the disclosure of the differences, is rather strong evidence 
of the general prevalence of such differences. 

(3) Although statistical investigation of animal behavior 
has been much less prosecuted than has either of the other 
types it is, nevertheless, within the limits of its availability, 
a very valuable method for revealing species differentia, its 
efficacy consisting in the fact that species may be compared 
with reference to different behavior traits taken one by one, 
and on the basis of quantitative data covering considerable 
samples of whole populations. The method is specially ap- 
plicable to the minute floating life of the seas and lakes, 
known as plankton, and is being much employed to this end 
at the Scripps Institution for Biological Research. It can- 
not be described in detail here, but consists essentially, as 
employed at this Institution, in collecting great quantities 
of organisms by agencies as nearly quantitatively constant 
and accurate as possible, in counting the organisms thus 
secured, and in correlating the biological values thus ob- 
tained with quantitative studies on the physical environment 
of the organisms, these environmental determinations being 
made simultaneously with the collection of the organisms. 
By this means one element in the behavior, that namely of 
the up-and-down journeys in the sea, long known to be per- 
formed by many oceanic species, has been studied with a fair 
degree of quantitative accuracy as to the extent of travel, 
time required for each journey, and environmental influence. 
A considerable series of species have been compared on this 

Organic Connection Between Physical and Psychical 281 

Two quotations are all that "space will permit us to give 
for showing what these investigations are bringing out on the 
subject of specificity in behavior. The animals referred to 
in the first quotation constitute a group of small, arrow- 
shaped worms known as the Chaetognatha bristle-jaws. 
''Each species occurring in the San Diego region has its own 
definite and specific manner of vertical distribution, just as 
truly as it has its own morphological characteristics." 22 

Similar results have been obtained by the same methods 
applied to a very different group of animals, minute crus- 
taceans of the ubiquitous order Copepoda. The investiga- 
tions on this group have been made by Dr. C. O. Esterly, 
and the results are specially interesting in this case, because 
Doctor Esterly has applied laboratory experimentation, to 
some extent, to the same animals, and has found a good de- 
gree of concordance in the results of the statistical and the 
experimental investigations. "A heterogeneous assortment of 
forms may be obtained in the same collection but each has its 
own way of reacting toward the elements of its environ- 
ment." 23 

It is the indubitable trend in one direction of the vast 
evidence from these three quite different classes of research 
on animal behavior that to my mind justifies such a con- 
ception of specificity of psychical and reactive animal life 
as that formulated above. Something of the probable mean- 
ing of this specificity we shall see in the next chapter. 


1. Loeb ('02) 189 13. Huxley, J. S 156 

2. Morgan, C. L 10 14. Darwin, C I, 353 

3. Royce 328 15. Kellogg, V. L 648 

4. Royce 329 16. Nature Lovers' Library II, 218 

5. Holmes ('11) 92 17. Nature Lovers' Library 

6. Holmes (11) 93 III 100 

7. Peckham, G. W. and E. S. 53 18. Seton I, 360 

8. Holmes ('11) 21 19. Groos 15 

9. Jennings ('06) 49 20. Wheeler ('10) 518 

10. Holmes ('11) 95 21. Ritter ('18) 119 

11. Wallace, A. R 555 22. Michael 46 

12. Huxley, J. S 151 23. Esterly 11 

Chapter XXIV 



Remarks on the Hypothetical Character of This Chapter 

HYPOTHESIS and theory will dominate in the task 
upon which we now enter and in this respect the 
present chapter will differ sharply from the preceding chap- 
ters. Fact, description, classification, and restrained gen- 
eralization have been the leading motives up to this point. 
One main and several subsidiary hypotheses will be central 
in the discussion. Into the presence of these will be sum- 
moned many of the facts and generalizations previously set 
forth. The purpose in this summoning will be on the one 
hand to test the hypotheses by the facts and generalizations 
and on the other hand to see how the facts will look in the 
light of the hypotheses. 

This announcement of the hypothetical and theoretical 
character of the task now before us, will give us two advan- 
tages : It will justify a dogmatic form of expression at times 
which we should not otherwise feel privileged to use ; and 
will justify a brevity of treatment which would not be pos- 
sible were we aiming at thorough generalization and demon- 
stration. Hence the justification of undertaking to deal 
with so vast and vital a subject in the limits of a sketch. 

The Natural History Method and the Study of One's Self 

Insistent as I have been on the importance of the natural 
history way of approaching the phenomena of the living 


Sketch of an Organismal Theory of Consciousness 883 

world, in entering upon the present discussion I must em- 
phasize this more than ever and must call attention to the 
particular character of this importance in our present un- 

The natural history method of viewing organic beings is 
per se the comprehensive method, one of its best mottos 
being, as we have repeatedly seen, "neglect nothing." That 
knowledge of organisms separates itself sharply into de- 
partments is no deterrent to the naturalist against utilizing 
any knowledge he may come upon that will contribute to his 
main aim that of understanding organisms. Who or what 
shall restrain me from observing and carefully thinking 
about any fact of my own being which promises to help me 
on my road to such understanding? The foremost zoolo- 
gists, of modern times especially, have amply recognized and 
freely used this principle so far as all physical and some of 
the lower psychical attributes are concerned. But w T hen it 
comes to man's higher psychical attributes, zoologists have 
usually said, sometimes expressly, sometimes tacitly, that 
these belong to a wholly different realm, a realm with which 
we have little or nothing to do. And their position of 
"hands off' as touching man's higher psychic life, has re- 
ceived the readier, fuller sanction in that it has accorded 
well with the prevalent views and practices of those students, 
anthropologists, economists, sociologists, and ethicists who 
have made these higher reaches of human life their special 
fields of inquiry. But the course of nature can not be per- 
manently thwarted. Such an attempt to wrench human life 
asunder is bound to fail finally. In the several subdivisions 
of biology, normal advance has tended to stay the wrenching 
process, comparative psychology being notable in this ten- 

The opposition to such organic disunion consistently 
maintained throughout this book reaches its culmination in 
these chapters on psychic integration. In what follows we 

284 The Unity of the Organism 

shall pass more freely than ever from one phase or aspect 
to another, over the entire gamut of psychic life both in the 
individual and in the animal kingdom. If facts of my own 
subjective life will serve my purpose, I shall be as free to 
requisition them as to requisition facts of any phase or as- 
pect of my objective life. If the ethical or esthetic or social 
attributes of the human animal will best illuminate a point, 
these shall be brought in with as little misgiving as will be 
anatomical or embryological or physiological or instinctive 

So great store do I lay on this catholicity of attitude 
toward psychic life, that I shall show by a single instance 
that at least a few other present-day zoologists have some- 
what similar feelings about the zoological character of psy- 
chical phenomena. Referring to the controversies which 
have inevitably arisen over the problem of instinct, W. M. 
Wheeler says that such controversy "is pardonable, at least 
to some extent, since the subject itself presents no less than 
four aspects, according as it is studied from the ethological, 
physiological, psychological or metaphysical points of view." 
"From the first two of these," the author continues, "in- 
stinct is open to objective biological study in the form of 
the 'instinct actions.' These may be studied by the physiol- 
ogist merely as a regularly coordinated series of movements 
depending on changes in the tissues and organs, and by the 
ethologist to the extent that they tend to bring the organism 
into effective relationship with its living and inorganic en- 
vironment. But that these movements have a deeper origin 
in psychological changes may be inferred on the basis of 
analogy from our own subjective experience which shows us 
our instincts arising as impulses and cravings, the so-called 
'instinct-feelings'; and these in turn yield abundant material 
for metaphysical and ethical speculation." 1 From the 
context of these sentences we may infer that Wheeler rec- 
ognizes that the four aspects mentioned under which the 

Sketch of an Organismal Theory of Consciousness 285 

subject of instincts presents itself, represent the same num- 
ber of valid departments of man's mental life. The point 
I wish to make is that although a zoologist may recognize 
without cavil that speculation on psychological, ethical, 
and metaphysical problems which arise in connection with 
instincts, are legitimate activities of man, and might prop- 
erly deny that it is incumbent upon him to do much specu- 
lating of this sort, yet it would be incumbent on him to take 
due cognizance of these speculative attributes of the human 
animal. A truly scientific zoology can not justify itself in 
issuing a manifesto to the effect that certain attributes pre- 
sented by some animals do not fall within its province. It 
may more or less constantly neglect or refuse on practical 
grounds, to deal with certain attributes ; but that is a very 
different matter from a formal declaration such as many 
present-day zoologists make, that with these attributes 
zoology has nothing to do. Such a declaration is self-stunt- 
ing, if not self-stultifying, in that it is a virtual self-inhi- 
bition by zoology of its own growth. 

These reflections may be terminated by defining the mo- 
tives and the mental attitude with which I approach the 
great problem of consciousness. I come to it not as a meta- 
physician, not as a psychologist, not as a physiologist, not 
even as an anthropologist, but as an anthropological zoolo- 
gist ; as a zoologist who in course of his regular professional 
work takes up the animal group of which he himself is a mem- 
ber, chancing as he does to possess among other attributes 
that of knowing his own life directly, that is, through sub- 
jective or self-conscious experience, as well as indirectly 
through objective experience. 

Approaching the problem of consciousness in such an at- 
titude and for such a motive, it is impossible to view it other- 
wise than as one aspect of the larger problem of life gener- 
ally. For while the psychologically and metaphysically im- 
portant question of whether consciousness is coextensive 

286 The Unity of the Organism 

with life need not be raised by the naturalist, the indubitable 
fact that at least a large sector of life is conscious ; in other 
words, the fact that consciousness is a part of life, he can 
not ignore if he is to deal with consciousness at all. For the 
naturalist, then, no hypothesis or theory of consciousness 
can be satisfactory which is not clearly and expressly em- 
bedded in and an essential part of an hypothesis or theory 
of life generally. Our central hypothesis, drafted in ac- 
cordance with these principles, may now be given. 

Formulation of the Central Hypothesis 

All the manifestations which in the aggregate we call 
Life, from those presented by the simplest plants to those 
of a consciously psychical nature presented by man and 
numerous other animals, result from the chemical reaction 
between the organism and the respiratory gases they take, 
oxygen being almost certainly the effective gas for nearly 
all animals. An essential implication of this proposition 
is that every living individual organism has the value, 
chemically speaking, of an elementary chemical substance. 

Let us be promptly explicit in recognizing the character 
of the two propositions contained in this hypothesis. They 
are manifestly chemical in large part, and a complete demon- 
stration of their truth is impossible without the aid of chem- 
ical research focus sed directly upon them. But though 
clearly chemical, equally clearly they go beyond- -far beyond 
-present chemical knowledge. To speak of a whole organ- 
ism as equivalent to a chemical element seems at first sight 
not only unwarranted by positive chemical knowledge, but 
opposed by such knowledge. Furthermore, the term "re- 
action" as used in the first proposition undoubtedly seems 
quite foreign to the technical meaning which chemistry has 
attached to the word. Indeed so remote to say the least, 
are these fundamental propositions of the hypothesis from 

Sketch of an Organismal Theory of Consciousness 287 


definite chemical knowledge, that if they are entitled to rank 
as constituting a legitimate scientific hypothesis, this must 
be on grounds other than those of present-day technical 
chemistry quite as much as on those of such chemistry. In 
attempting, consequently, to establish the propositions on a 
true and useful hypothetical basis, it will be permissible to 
notice these other grounds first. 

Preliminary Justification of the Hypothesis as Such 

The proposition that each living individual has the chem- 
ical value of an elementary substance, will receive attention 
first, and the initial step will be to inquire what, in general, 
the criterion is of an elementary chemical substance. Here, 
for instance, is a lump of phosphorus. In virtue of what is 
it declared to be such a substance? Not primarily, let us 
specially notice, because the phosphorus is simple, that is 
to say, is an element in the sense of not being reducible to 
still simpler substances. Rather the basal criterion of its 
being a chemical substance is that upon its being brought 
into contact under certain conditions with certain other 
chemical substances, oxygen for instance, there is produced 
a third substance having very different attributes from either 
of the original substances. Transformation of substances 
chiefly through interaction upon one another is the founda- 
tion fact which has brought it to pass that substances are 
described as chemical. That is the fact upon which the 
science of chemistry primarily rests. Facts and problems 
of simplicity and complexity, relative and absolute, are 
later and secondary. The task of chemistry "consists in the 
investigation of substances and those of their processes by 
which the physical attributes of the substances undergo 
permanent changes." (Handzvorterbuch der Naturwissen^ 


Every adequate definition of chemistry and chemical sub- 

288 The Unity of the Organism 

stance and chemical action contains the idea of transforma- 
tion in one form or another. Clearness on this point is in- 
dispensable to our purpose. Chemistry is too often defined, 
even in elementary text books and in dictionaries, as thougli 
the "composition of matter" were its initial and most es- 
sential function. But this conception is surely contrary to 
the history and most essential nature of the science. There 
is, it seems, entire agreement among competent writers that 
scientific chemistry is a direct* descendant of Alchemy, and 
a very imperfect knowledge of the history of Alchemy re- 
veals the fact that the every-where present, normal trans- 
formations in nature, particularly in inorganic nature, were 
the foundation phenomena of this old art. One has only to 
recall the place held by the idea of the transmutation of 
metals, this idea having usually the practical aim of chang- 
ing the "base metals" into "noble metals." The "phil- 
osopher's stone" and the "great elixir" were magical some- 
things by which the transmutations could be accomplished. 

Greatly significant from our standpoint is the fact that 
one of the objectives of Robert Boyle (middle of the seven- 
teenth century), who, perhaps as much as any one man, is 
entitled to be called the father of experimental chemistry, 
was to rectify the false and mystical notions prevalent in his 
time about "Elements," "Principles," "Essences," etc. "Tell 
me what you mean by your Principles and your Elements," 
Boyle demanded, "then I can discuss them with you as work- 
ing instruments for advancing knowledge." 

What is "behind" the transformations forces, elements, 
principles, essences, spirits or what not- -is indeed an impor- 
tant and, properly asked, a legitimate question. But- -and 
here is the most vital fact of all it is a question which can 
not be raised even, until after the transformations have been 
observed, nor can an answer of objective value be given un- 
less the whole round of observed phenomena, the substances 
previous to transformation, the transformatory processes, 

Sketch of an Organismal Theory of Consciousness 289 

and the new substances, be accepted at their face value, that 
is to say, at a value which is as near to ultimate truth as any 
truth whatever, connected with the phenomena. 

The elemental constitution of bodies is an inference, al- 
ways and solely, drawn from their observed corporeal attri- 
butes. And chemistry is the science which assumes the task 
of drawing, elaborating, and systematizing these inferences 
on the basis of the transformation of the attributes. The 
meaning of the statement that chemistry is one of the natural 
sciences is that chemistry is the science which uses its natural 
history observations to penetrate still more deeply into the 
constitution of bodies. Natura a natwra vincitur, nature is 
surrounded by, is contained in nature, is as fundamental a 
truth for chemistry as for any other natural science. A 
living being is as much a natural body as is a piece of phos- 
phorus, and its obvious attributes, its outer-layer attributes, 
are as essential to its nature as are its inner, its hidden 
attributes. So any genuinely transformatory changes, and 
genuinely new products arising through the reaction between 
the living body and some other body is so far chemical in 
nature, and the reacting bodies are so far chemical. 

A long step toward justifying the proposition that each 
individual living organism has the value, chemically, of an 
elementary substance, will be taken if it can be shown that 
any qualitatively new product whatever results from the 
interaction between the organism acting as a unit, as one, 
as an element, and some other element. Having regard to 
the entire world of living beings, the chances for finding new 
products which may have arisen in this manner are prac- 
tically if not theoretically infinite. Manifestly, then, only a 
very small sector of the entire range of such possible produc- 
tions can be searched. It must, consequently, be our aim, as 
always in handling inductive natural history evidence, to 
choose for examination evidence which shall be most clear- 
cut, most illustrative, and most convincing. 

290 The Unity of the Organism 

The sector of organic phenomena best capable of yielding 
such evidence is, I believe, exactly this of psychic life. And 
within the great range of this life, the higher conscious life 
of man is most replete with the evidence we seek. Again 
within the range of man's higher life, each individual's own 
private life, even his subjective life, his consciousness, is the 
evidence most certain and convincing. Translating this last 
statement into familiar language, one sees that it is only 
another way- -the scientific way of affirming the truth, that 
the greatest of all certainties of which man is capable is that 
of his own existence. I am saying, virtually, that when we 
analyze, after the manner of objective science, this old fa- 
miliar affirmation about certainty, and carry the analysis 
as far as we are at present able to, we find that the sense, or 
better, the feeling of certainty of self-existence and self- 
identity is in last analysis one of the effects of a transforma- 
tory interaction between ourselves and some substance (oxy- 
gen?) in our breath, as stated in the first of our two propo- 

That proposition seems then to be hardly more than a 
recognition that psychic phenomena containing at least the 
germ of consciousness is a kind of chemical product which 
has not heretofore been clearly recognized as such, the lack 
of recognition being due to the strangeness of the product as 
compared with any chemical products with which experimen- 
tal chemistry has hitherto occupied itself. But looked at in 
a really broad and deep way, is it any more difficult for me 
to interpret a state of consciousness in myself to be a result 
of chemical action between me and the air (oxygen?) I 
breathe, than for me to interpret the dim greenish-white 
luminosity of a piece of phosphorus to be a result of the 
chemical action between the phosphorus and the air essential 
to the glowing? From a purely chemical standpoint I do 
not believe we have any ground for holding that some prod- 
ucts of chemical reaction are more comprehensible or less 

Sketch of an Organismal Theory of Consciousness 291 

comprehensible than are others. 

Chemically viewed the problem now on our hands is en- 
tirely one of fact fact as determined by observation alone, 
and by observation with the aid of experimentation. If it 
can be shown that each individual conscious being really does 
behave like a chemical substance in the process of reacting; 
and if the result of such reaction can be shown to have even 
one of the essential marks of a chemical product, both propo- 
sitions of my two-parted hypothesis are warrantable and the 
hypothesis becomes genuinely scientific a genuine "working 
hypothesis" one, that is, for bio-chemistry to take seri- 

More Systematic Justification of the Hypothesis 

That the propositions are demonstrable to the extent of 
the demand just indicated is my contention. This conten- 
tion I will now try to make good and will begin with a few 
remarks on a question concerning the hypothesis which 
ought to arise instinctively in the mind of every one. That 
question is : Does such a conception of psychic life and 1 con- 
sciousness as that contained in our hypothesis imply any real 
infringement upon or derogation from me, in the deepest 
sense a real entity properly designated by the terms person 
and personality? 

On saying that this query ought to arise mstinctively, I 
do not mean ought in the ethical sense, but in the organismal 
sense. That is, in a sense which implies that the very nature 
of the conscious organism is that it is not only self-existent 
in a measure like every natural object, but that it is self-iden- 
tifiable, and within certain bounds, self-determinative of its 
own acts. Now recognizing it to be thus by its "very na- 
ture" is only another way of recognizing that it is so in its 
instincts as well as in its physical organization. But since 
instinct is more fundamental, more deep-rooted in the or- 

The Unity of the Organism 

ganism than is intellect, as phylogenic and ontogenic psy- 
chology make clear, if a pronouncement implying a de- 
rogation from the reality and natural prerogatives of the 
individual be issued from the intellect, a response of protest 
and antagonism would be expected from instinct. This would 
be expected as an ordinary organic impulse to self-defense 
and self-preservation. 

The Nature of "Outer" or Objective and "Inner" or 


What we have to do consequently is to scrutinize the con- 
scious individual in order to see if it presents any uniqueness 
of attributes and of transformatory power in reacting with 
other bodies that is on a par with the uniqueness of an ordi- 
nary chemical substance in the same respects. Now it is, 
as suggested some pages back, exactly in the conscious, the 
subjective life, that such uniqueness is most easily demon- 
strable. There are several ways in which the conscious indi- 
vidual manifests this uniqueness. A particularly convincing 
way, I think, is in the relation between what are commonly 
known as the objective, or "outer," and the subjective or 
"inner" sides of mental life. This, consequently, will be the 
approach to the subject chosen by us and we will enter upon 
it by returning to Royce, first to his "Outlines of Psychol- 
ogy," then a little later to some of his specifically philosophi- 
cal writings. 

In the first chapter of the Outlines, devoted to initial defi- 
nitions and explanations, Royce states, simply and clearly, 
a distinction "between our physical and mental life," which 
elsewhere he has worked out with great elaboration. Thus: 
"Physical facts are usually conceived as 'public property,' 
patent to all properly equipped observers. All such observ- 
ers, according to our customary view, see the same physical 
facts. But psychical facts are essentially 'private property,' 

Sketch of an Organismal Theory of Consciousness 293 

existent for one alone. This constitutes the very conception 
of the difference between 'inner' psychical or mental, and 
physical or 'outer' facts." 4 

Ever-present, and obvious as is the comparison here made, 
it nevertheless is of so great importance that we must stop 
and reflect upon it, for we shall surely fail to grasp the full 
measure of what is to follow if we are lukewarm toward one 
of the elements of it. The element I refer to is the unique- 
ness, the essentially personal character of inner as contrast- 
ed with outer facts. Every normal person is ready enough 
to insist that his thoughts, his feelings, his emotions and all 
the rest of his higher psychical experiences are his and his 
alone. The tremendous reality and force of the rights of 
"private opinion," of "personal conscience" and so forth, 
among civilized men, hardly need to be expatiated on. 

The character of the uniqueness of these experiences, how- 
ever, concerns practical living less vitally, so we give it less 
attention. The whole vast range of my mental life, from 
the lowest, simplest, vaguest sensations to the highest, most 
bewildering complex emotions, passions, imaginings and 
thoughts, are my own, absolutely, so far as other persons 
are concerned. I cannot share them to the least extent with 
another person. Of course I can let others, especially my 
most intimate associates, my dearest friends, know a good 
deal about these experiences of mine. But after all, gladly 
as I would share many of them with these friends, it is utter- 
ly impossible for me to do so. My experiences must remain 
wholly outside of their consciousness. No two persons can 
have the same experience any more than they can have the 
same hands or stomachs. Nor is this all. If mental life is 
subject to the general biological laws of variation into which 
we have latterly gained much insight, I am obliged to sup- 
pose that these experiences of mine, the whole retinue of sen- 
sations, feelings, emotions and thoughts, differ somewhat 
from the corresponding experiences of other persons. And 

294 The Unity of the Organism 

all observation confirms this supposition much of it strong- 
ly. Inferential evidence could hardly be stronger than that 
my particular emotional response to opera singing, for 
example, is quite different from that of many other persons. 
Obviously we are here skirting the edge of what modern 
realism in formal philosophy calls pluralism, and deals with 
in part as the question of whether percepts are strictly indi- 
vidual and personal. No philosopher with whose views I 
have become acquainted, has discussed this question so fully, 
and in my opinion, so illuminatingly as Sellars. The follow- 
ing sentences taken from his chapter, The Advance of the 
Personal, show clearly, it seems to me, that the conclusions 
he has reached, working from the purely philosophical side, 
are essentially the same as those arrived at by me, advancing 
from the biological side : "What may be called the sensory 
content of our percepts is important,--! do not wish to 
be understood to belittle it, but so are the meanings which 
arise in conection with our bodily activities and motor ad- 
justments to stimuli. Here again, we are face to face with 
individual factors in perception which even the idealist must 
recognise and somehow explain. Evidently, perception is not 
a mere passive presentation, but a construction whose gene- 
tic elements can be partially traced. Finally, let us call to 
mind that percepts are continuous with feelings and with the 
so-called organic sensations. . . . Once vaguely objective, 
feeling is now considered subjective or personal." Many 
other sentences and paragraphs of like purport could be 
quoted from this author. I have selected this for the two- 
fold reason that it indicates the measure of my agreement 
with his view as to the personal character of percepts and 
the rest of conscious life; and at the same time indicates 
wherein I shall have to out-do him in the matter of validat- 
ing the individual. A part of our task, to be reached a little 
later, will be to show that although feeling and all the rest 
of psychic life is indeed subjectively personal, it is also 

Sketch of an Organismal Theory of Consciousness 95 

objectively personal. In other words, it will be my task to 
remove, or at least to show the way to remove, the vagueness 
which Sellars asserts, rightly, has hitherto clouded this 
side of personality. To do this thing is, indeed, one of my 
most important chances to contribute to a "better philosophy 
of life." 

But since our psychical life, especially our conscious life, 
is a vast incalculably vast complex of experiences, of 
"contents," sounds, sights, memories, feelings, ideas, many 
of which are set off very sharply from the rest, are clearly 
characterizable, and are wonderfully persistent; and since 
innumerable of these are coming along all the while which 
have much of genuine newness about them ; and since further, 
these contents of consciousness are intertwined with and are 
determinative of a vast complex of other contents called voli- 
tions which in turn are linked up with and are more or less 
directive of bodily activities of many kinds, some purely re- 
flex and some instinctive, it seems impossible to escape recog- 
nizing, even if one wanted to, that if the verb "to create" has 
any definite meaning at all the normal, self-conscious animal 
organism is about the most creative thing we know or can 
conceive. Indeed it is altogether likely that the very notion 
of creation, whether natural or supernatural, came initially 
from the creative activity and the impulse to such activity, 
of man himself. 

We may justly say, I think, that we know all creativeness, 
chemical creativeness with the rest, through being in our own 
deepest natures creative, that is, transformative and trans- 
formative in the way which we call chemical. We learn 
about the processes of life and call some of the most essen- 
tial of them chemical just by performing those processes as 
some of our most essential attributes. A portion of the pro- 
cess which goes on within us, together with the corresponding 
product, constitutes what we call the science of bio-chemis- 
try. This means that according to our hypothesis "objec- 

296 The Unity of the Organism 

tive" and "subjective," or "outer" and "inner" as applied to 
life, are something quite different from what they have been 
either in traditional philosophies, or in most, at least, of 
recent psychology. "When we speak," Royce writes, "of 
our physiological processes as internal, the word 'internal,' 
although it here generally implies 'hidden, in whole or in part, 
from actual outer observation', does not imply 'directly felt 
by us ourselves.' My hypothesis implies a denial of the 

correctness of this statement. I say that in the sum total 
of the "contents of consciousness," a nether segment, as one 
might call it, of physiological processes is "directly felt by 
ourselves." There is no content of consciousness which does 
not contain an element that is internal or subjective in what- 
ever sense any other content of consciousness is internal or 
subjective. And per contra, there is no content of conscious- 
ness which is not objective to some extent, in whatever sense 
any other content of consciousness is objective. The mind, 
according to this conception, is not something which uses 
the brain or any other part of the organism merely as a tool 
with which to make thoughts and other contents of con- 
sciousness. Nor on the other hand is consciousness of the 
nature of a secretion, the gland for which is the brain, though 
unquestionably the brain has an essential part in the pro- 
duction of thought and the higher contents of consciousness. 
Among the consequences of the reaction between the or- 
ganism and the air we breathe are consciousness with its 
marvellously rich and varied contents. 

But at this point I must specially request the reader to 
notice that I am not pretending to describe and explain all 
the contents of consciousness. In other words it is not a 
theory of knowledge, but a theory of consciousness that I 
am sketching; and knowledge in the strict sense, and con- 
sciousness are very different. They differ, according to my 
understanding, much as the fully developed, physical organ- 
ism differs from the living substance, or protoplasm, of 

Sketch of an Organismal Theory of Consciousness 97 

which the organism is composed. Consequently I am not 
even concerned primarily with sensation in so far as this im- 
plies sense organs or even nerves and nerve terminals of the 
simplest kind. Rather I am dealing with the stages and con- 
ditions antecedent to consciousness and in which it is latent, 
in much such way as the cytologist when he studies the living 
substance of all sorts of tissue-cells is not dealing with organs 
and the organism in the full sense, but only with their sub- 
strata. But although it is not knowledge, properly speaking, 
either in its conceptual or perceptual aspect that I am dis- 
cussing, since my enterprise does take me across the border 
line and a short distance into the realm of knowledge, I must, 
in the interest of historical continuity and setting, say. a 
little more than I have said about the general nature of 

My assertion should be taken literally that there is no 
content of consciousness which is purely either subjective or 
objective, inner or outer, conceptive or perceptive, ideational 
or impressional, or whatever form of expression be given the 
antithesis here implied. That every content of consciousness 
which exists or can be conceived has an essential element of 
both members of the antithesis is exactly what I mean. To 
illustrate, even the axioms, postulates, or whatever else may 
be counted as most ultimate in mathematics contain an ele- 
ment of the outer, or objective, as well as of the inner, or 
subjective. These mathematical contents of consciousness I 
single out to illustrate my meaning because they have been 
clung to by philosophers and scientists more tenaciously than 
any others as purely subjective or mental. And further 
there is a strategic gain in this reference to mathematics in 
that it brings into the open the fundamental opposition of 
my hypothesis to one main root of Cartesian philosophy; 
the philosophy, that is, from which the modern doctrine of 
psycho-physical parallelism has grown. Our thinking, which 
Descartes held proves our existence, really proves it only in 

298 The Unity of the Organism 

so far as it shows that among the activities essential to the 
human organism thinking is one. In other words the "there- 
fore" in "I think, therefore I am," is true only because "I 
am, therefore I think," the reverse proposition, is also true 
and includes the other truth. The lesser truth is true be- 
cause it is an essential part of the larger truth, much in 
the same way that the cells of a multicellular organism are 
alive because they are essential parts of the organism. 

We need not inquire how, from this serious shortcoming of 
Descartes' description of psychic life Descartes went on to 
the conclusion that "there is nothing really existing apart 
from our thought" and that "neither extension, nor figure, 
nor local motion, nor anything similar that can be attrib- 
uted to body, pertains to our nature, and nothing save 
thought alone ; and, consequently, that the notion we have of 
our mind precedes that of any corporeal thing, and is more 
certain, seeing we still doubt whether there is any body in 
existence, while we readily perceive that we think." 18 Nor 
need we concern ourselves with the voluminous and tedious 
reasonings by which a considerable number of moderns, fol- 
lowing Descartes's lead, have convinced themselves that they 
have "reduced" all reality or at least all reality that really 
amounts to anything, to quantity. Enough now to remark 
that every modern biologist who really accepts the basal data 
of his science, must agree that "Psycho-physical paralellism 
. . . stands to-day as the scandalous but irrefutable conse- 
quence of postulating a material world without qualities and 
a world of minds that lack spatiality and exists nowhere." 
One way of characterizing my hypothesis would be to say 
that it is an effort to remove this scandal by showing where- 
in the postulation noted by Dr. Montague is not true. 

The genetic relationships of my hypothesis can be still 
farther indicated by coming on down from Descartes to 
Hume then from Hume to Huxley and finally to G. F. Stout 
and John Dewey as philosophers of to-day. Hume's nom- 

Sketch of an Organismal Theory of Consciousness 299 

enclature for the subjective and objective sides of man's 
psychic life is "Relations of Ideas" for the first, and "Mat- 
ters of Fact' : for the second. Of the first kind says Hume, 
"Are the sciences of Geometry, Algebra and Arithmetic ; and 
in short, every affirmation which is intuitively or demonstra- 
tively certain." . . . "That three* times five is equal to half 
of thirty, 9 ' is a simple illustration of the relation of 
ideas. And, "Propositions of this kind are dis- 
coverable by the mere operation of thought, with- 
out dependence on what is anywhere existent in the uni- 
verse." 20 And further on, Part 2, same section, we read: 
"It must certainly be allowed, that nature has kept us at a 
great distance from all her secrets, and has afforded us only 
the knowledge of a few superficial qualities of objects; while 
she conceals from us those powers and principles on which 
the influence of those objects entirely depends." Then Hume 
goes into a discussion of the operations and relations of the 
"superficial qualities" and "secret' 1 powers of objects which 
is so similar to my treatment of the relation of the organism 
to the attributes of certain objects (chapters 20 and 21 this 
book, and, more particularly, my essay Is Nature Infinite? 21 ) 
that it seems as though his words must have been in my mind 
when I thought out what I have there written, though I cer- 
tainly was not conscious of Hume's views. And this sub- 
conscious influence appears the more probable in that I have 
almost conclusive proof of having read his argument not long 
before my own was written. I am certain, however, that if 
his statements were in my mind they were only in its pro- 
conscious part and were not nor ever had been in its full- 
conscious part. In other words, if I had read his words I 
had not grasped their full significance. This probable in- 
stance of the "sub-' or "pro"-conscious I refer to not so 
much because of its interest in this instance, as because of its 
bearing on my conception of the nature of consciousness. 
The discussion by Hume to which I refer is that in which he 

300 The Unity of the Organism 

talks about the sensible qualities and the "secret powers" 
of the bread we eat. "Our senses inform us of the color, 
weight, and consistence of the bread," he says, "but neither 
sense nor reason can ever inform us of those qualities which 
fit it for the nourishment and support of a human body." 
The particular puzzle upon which Hume comes in this 
matter is the fact that although the examination here and 
now of a natural object gives us absolutely no clue as to what 
latent attributes ("secret powers," he calls them) the ob- 
ject may possess, when we examine a second object of the 
same kind we assume that the same secret powers are pos- 
sessed by the second object. "If a body of like colour and 
consistence with that bread, which we have formerly eat, 
be presented to us, we make no scruple of repeating the ex- 
periment, and foresee, with certainty, like nourishment and 
support. Now this is a process of the mind, of thought," 
Hume goes on to say, "of which I would willingly know the 
foundation." "The bread," he says, a little farther on, 
"which formerly I eat, nourished me ; that is, a body of such 
sensible qualities was, at that time, endued with such secret 
powers : but does it follow that other bread must also nour- 
ish me at another time, and that like sensible qualities must 
always be attended with like secret powers? The conse- 
quences seem nowise necessary. At least, it must be acknowl- 
edged that there is here a consequence drawn by the mind; 
that there is a certain step taken; a process of thought, 
and an inference, which wants to be explained." Then after 
a little further argument to show the necessity of recog- 
nizing such a process we find this to me exceedingly interest- 
ing passage: "There is required a medium, which may en- 
able the mind to draw such an inference, if indeed it be drawn 
by reasoning and argument. What that medium is, I must 
confess, passes my comprehension; and it is incumbent on 
those to produce it, who assert that it really exists, and is 
the origin of all our conclusions concerning matter of fact." 

Sketch of an Organismal Theory of Consciousness 301 

The great merit here shown by Hume is his ability to push 
the analysis of his problem to the very limit of the positive 
information he had to go on, recognise exactly wherein his 
information was lacking, and then stop without running off 
into a purely speculative substitute for his deficient knowl- 
edge. According to my hypothesis the unknown "medium" 
which he saw must exist, the researches of a century and a 
half since he wrote, in chemistry, physiology, general zoology 
and botany, and psychology, have enabled us to see is the 
individual animal organism reaching with the respiratory 
substance (oxygen?) it takes in. In this one particular and, 
from the standpoint to which we have been accustomed, very 
peculiar case, the reaction is at one and the same time part 
of the essence of both ideas and impressions in the Humean 
sense, the reaction being the "medium" or the "certain step" 
by which the inference is drawn, this inferring being possible 
because of the continuity of the organism as a person, or 
self, and the persistence of the respiratory substance as the 
same identical thing from the past through the present into 
the future. 

We will now notice how Huxley, because of his much more 
extensive knowledge of the structure and function of animals 
than Hume possessed, was able to draw still closer than Hume 
could to the heart of the old Mind-Body puzzle. The gist 
of Huxley's position on, and contribution to, the problem 
can conveniently be presented through his remarks on the 
question of innateness of various aspects of psychic life, 
these remarks occurring in his essay on Hume. After point- 
ing out that neither Locke nor Hume seemed to know exact- 
ly what Descartes, the originator of the modern conception 
of innate ideas, meant bv his phrase "idees naturelles," Hux- 

^ 9 

ley quotes Descartes as follows: 'I have used this term in 
the same sense as when we say that generosity is innate in 
certain families; or that certain maladies such as gout or 
gravel, are innate in others ; not that children born in these 

302 The Unity of the Organism 

families are troubled with such diseases in their mother's 
womb; but because they are born with the disposition or 
faculty of contracting them." Then after further quota- 
tions to the same effect Huxley writes : "Whoever denies 
what is, in fact, an inconceivable proposition, that sensations 
pass, as such, from the external world into the mind, must 
admit the conclusion here laid down by Descartes, that, 
strictly speaking, sensations, and a fortiori, all the other 
contents of the mind, are innate. Or, to state the matter in 
accordance with views previously expounded, that they are 
products of the inherent properties of the thinking organ, in 
which they lie potentially, before they are called into exist- 
ence by their appropriate causes." 

The upshot of this clearly is that innate for Descartes and 
Huxley means hardly anything else than hereditary, as ap- 
plied to the psychical as well as to the physical attributes of 
animals. The ample justification in our day of the view 
that psychical attributes are hereditary should, it would 
seem, restore to full standing in biology, the conception of 
innate ideas only, of course, in a very different sense from 
that into which later Idealists have perverted it. 

It is in this discussion that Huxley makes one of the most 
direct and unanswerable arguments against materialism that 
can be made : "The more completely the materialistic posi- 
tion is admitted, the easier it is to show that the idealistic 
position is unassailable, if the idealist confines himself with- 
in the limits of positive knowledge." That is to say, if the 
materialist insists that all traces of innateness of ideas and 
other contents -of the mind must be repudiated, he virtually 
contends that heredity of whatever sort, whether of physical 
or psychical attributes, must be repudiated. With this con- 
ception of innateness in the entire psychic aspect of the 
organism before him Huxley asks : "What is meant by ex- 
perience ?" 

"It is the conversion," he replies, "by unknown causes, of 

Sketch of an Organismal Theory of Consciousness 303 

these innate potentialities into actual experiences." 24 Now 
these "unknown causes" are, according to my view, essential- 
ly the same as the "medium" which Hume recognized must 
exist for making the "step" possible from the "superficial 
qualities" to the "secret powers" of natural objects and from 
the "secret powers" of one object to those of another. They 
are, to repeat, the reaction of the organism in its latently 
psychical aspect, with "the breath of life," that is, with the 
oxygen, or whatever be the gaseous constituent of the air 
which is active in respiration. And I believe we can see to 
a considerable extent why Huxley considered these causes as 
wholly unknown. It was because physiology and bio-chemis- 
try in his day were not yet able to view the organism from 
the standpoint of physical chemistry. Because of this ina- 
bility Huxley nor any other physiologist of his period had 
an adequate structural ground-work for thinking organis- 
mally about living things. They were consequently obliged, 
really, to think of all psychic phenomena, and consciousness 
with the rest, as being restricted to the nervous system. 
That such was Huxley's view at any rate, we know from 
his own words : "No one who is cognisant of the facts of the 
case nowadays doubts," he writes, "that the roots of psychol- 
ogy lie in the physiology of the nervous system." The im- 
portant revision of this statement which our hypothesis calls 
for is that while the roots of psychology are indeed in the 
nervous system they are by no means in that system alone. 
They pass through it to a much deeper level, so to speak, and 
in passing draw great nutriment from it. 

In a brief but important paper starting off with the prop- 
osition that a philosopher can not legitimately question the 
existence of the external world- -that all he can rightly do is 
to inquire what that world is and how we can know it at all, 
G. F. Stout comes to the kernel of the problem in considera- 
bly the same way that Hume and Huxley came to it. "For 
primitive consciousness and for our own unreflective con- 

304 The Unity of the Organism 

sciousness," he says, "sense experience and the correlative 
agency which conditions it coalesce in one unanalysed total 
object. They coalesce in such a way that the sense-presenta- 
tion appears as possessing the independence of the not-self, 
and the independent not-self seems to be given with the same 
immediacy as the sense-presentation." And, "this complex 
but unanalysed cognition," Stout continues, "is the germ 
from which our detailed knowledge of matter develops." 38 
If proved true my hypothesis would be a considerable for- 
ward step, I believe, in analysing this "unanalysed cogni- 
tion." For although Stout's assertion "the independent not- 
self is not matter" seems at first sight to exclude oxygen or 
any other constituent of our breath from such a place in 
the external world of his conception as that which it has in 
that world according to my conception this exclusion is, I 
think, only seemingly so, for a sentence farther on the author 
says matter "essentially includes the qualification of the in- 
dependent not-self by the content of sense-experience." The 
seeming discrepancy is probably due to the generality of the 
term matter. I too would say that the "independent not- 
self" is not matter were I to mean by matter the total sub- 
stance of the external world. But in the sense that the effec- 
tive respiratory gas (oxygen supposedly) is matter, my 
hypothesis would require me to hold that the not-self has an 
essential material component, which component is really the 
attribute of the gas in virtue of which it reacts with the 
organism in the peculiar way it does to produce conscious- 
ness. It seems to me that what Stout seeks in the "quali- 
fication of the independent not-self by the content of sense- 
experience" is the immediately consciousness-producing attri- 
bute of the respiratory gas. We might state the point this 
way: Oxygen (or the effective respiratory gas) has a double 
status in human consciousness. First and most fundamental- 
ly, it has the status of an immediate and essential participant 
in producing all consciousness whatever; and second it has 

Sketch of an Organismal Theory of Consciousness 305 

the status of an indirect participant in producing the par- 
ticular consciousness which we call observational knowledge 
of the gas. Our knowledge of this one gas is due to two 
things, (1) to our reaction to it through our sense organs 
in the usual psychological meaning of react; and (2) to our 
reaction with it through the protoplasmic basis of all con- 
sciousness, reaction in this case having the meaning which 
chemistry has given the word. What the relation is be- 
tween the attributes of the gas in virtue of which it reacts 
with the organism in these two ways, and also what the rela- 
tion is between the attributes of the organism in virtue of 
which it reacts with the gas in these two ways, are questions 
with which a theory of knowledge would deal but which lies 
outside of the scope of this sketch, which, as has already 
been said, restricts itself to a theory of consciousness. I 
may, however, refer in passing to the fact that chemistry 
appears to be all at sea on the problem of the relation be- 
tween the chemical and the physical attributes of all sub- 
stances whatever ; so the difficulties about oxygen in this one 
particular are not an unshared difficulty. 

Finally, to bring this exposition of the historical setting 
of my hypothesis down to the present hour, I call attention 
to the way the hypothesis connects with the best that formal 
philosophy in our own day has done, or as I suspect is 
competent to do, towards making out what "experience" is. 
No philosopher with whom I have met has gone farther in 
this direction than John Dewey. In his recent essay, A Re- 
covery of Philosophy, we read : "Dialectic developments of 
the notion of self-preservation, of the conatus essendi, often 
ignore all the important facts of the actual process. They 
argue as if self-control, self-development, went on directly as 
a sort of unrolling push from within. But life endures only 
in virtue of the support of the environment. 9 ' The italics 
are mine and mark the most vital part of the quotation for 
us. And a page farther on: "Experience is no slipping 

306 The Unity of the Organism 

along in a path fixed by inner consciousness. Private con- 
sciousness is an incidental outcome of experience of a vitally 
objective sort; it is not its source. Undergoing, however, 
is never mere passivity. The most patient patient is more 
than a receptor. He is also cm agent a reactor." . . . 
Again the italics are mine. I take the liberty to end the quo- 
tation at "reactor" though the remaining part of the sen- 
tence is important for Dewey's particular purpose. But my 
aim is different. I want to fix attention on the two state- 
ments italicised for the purpose of showing how my hypo- 
thesis connects with Dewey's general conception of experi- 
ence. When Dewey says life endures only as supported by 
the environment, he is speaking in very general terms, having 
reference, I imagine, more to social and other bulk aspects' 
of environment. My hypothesis, on the contrary, makes the 
dependence of life on environment exceedingly specific in that 
it undertakes to show the particular thing in the environ- 
ment, namely, the respiratory part of the atmosphere, which 
is physiologically basal to self-development and self-pre- 
servation. The Self which traditional philosophy has strug- 
gled so hard to understand is literally, the human organism, 
according to my hypothesis. And when in this discussion I 
speak of it as reacting with the respiratory air to produce 
consciousness, I am using the verb to react in a very specific, 
physico-chemico-biological sense, while Dewey is using it in 
a general sense, and explicitly at least, with only a psy- 
chological implication. 

The "self : which I am suggesting does indeed imply 
"another" no less unequivocally than does the "self" of ad- 
vanced social psychology. But the "self' ; and the "other" 
implied by my hypothesis differ from those of current philo- 
sophical theory in that the roots of both are not only in 
the social relationships of the human species, but extend 
right on through these into sub-human relationships, even 
down into the very constitution of inorganic nature. The 

Sketch of an Organismal Theory of Consciousness 307 

"self" and the "other" of my conception are more personally 
objective, and more cosmic in their affinities, than are the 
"self" and the "other" of social psychology. 

Continuing now with our examination of the foundation 
of my hypothesis I find it convenient, especially because of 
my reference a few pages back, to Huxley's unanswerable 
contention for an essence of truth in both materialism and 
idealism, to call attention to a natural history fact in the 
higher mental life of man which I take to be a strong con- 
firmation of the contention. This fact concerns the general 
difference between what are commonly known as the mate- 
rialistic and the idealistic attitudes of mind. This difference 
comes, I believe, to the same thing finally, as the difference 
between the objective and subjective attitudes, and is also 
the difference, at bottom, between what in rather loose though 
prevalent expression, is called the difference between the 
scientific and the philosophic attitudes. It would seem that 
the philosopher who declares himself to be an Absolute Ideal- 
ist, as Royce does, is under heavy obligation, especially if 
he enters the field of psychology, to explain the fact that the 
originators of great interpretative ideas of nature have in- 
variably recognized that their hypotheses must be "proved" ; 
that is, that the subjective experience which constitutes the 
hypothesis must be found to have its counterpart in the ex- 
ternal world of sense. If "Reason creates the world," even 
in the recondite meaning of Royce's philosophy, how hap- 
pened it that Newton should have been so "restless" for evi- 
dence of an objective, an external counterpart to the subjec- 
tive result he had reached by mathematical reasoning, that 
he held back his reasoned creation for sixteen years, waiting 
for the proof, the sense-perceptual or at least the sense- 
perceptible experience, that should round out his reasoned 
truth? May not, I ask, the very kernel of the difference 
between science at its best and philosophy at its best be in 
this, that the typical scientist is somewhat deficient in "rest- 

308 The Unity of the Organism 

lessness," adopting Royce's terminology, for internal or sub- 
jective reality; while the philosopher of the schools is some- 
what deficient in restlessness for external or objective reality? 
We could say with almost literal chemical accuracy that the 
curiosity and eagerness of the naturalist for yet unobserved 
objective truth is due to an unsatisfied affinity which is weak, 
or in some instances, wholly lacking, in the subjective idealist. 
The facts which seem to justify our chemico-organismal 
hypothesis of conscious psychic life, seem also to imply a 
complete interpenetration of objective science and idealistic 

As to the Lowest Terms of Self -Consciousness 

Let us now veer our course in examining self-conscious life, 
and see what can be made out about its roots and rootlets 
instead of about its fruitage. 

We are often reminded that our knowledge about our in- 
ternal organs, our heart, liver, lungs, et cetera, comes only 
through observations by the anatomist and physiologist; 
that we are quite unconscious of these organs in our own 
bodies, especially if they are working normally. Now I 
point out that to be perceptually conscious of a liver, let us 
say, as a specialized morphological entity performing its 
appropriate functions, is a very different matter from being 
conscious of those primal, undifferentiated processes which 
are basal to life itself, and so are common to all the tissues 
whether liver, muscle, brain, or what not, so long as they are 
actually living. That that which is truly organic, in the 
sense of pertaining to the fully constituted organism, must 
be regarded from this standpoint as well as from the stand- 
point of their final state of differentiation, is one of the 
common-places of modern biology. Let a person in as near- 
ly perfect health as he ever experiences, do his best to elimi- 
nate all external and internal stimuli of his specialized 

Sketch of an Organismal Theory of Consciousness 309 

sensory parts ; also all remembering, all feeling of the usual 
kind, all imagining, and all thinking. Then let him answer 
the question: How do I know I am alive? An undertaking 
of this sort is wholly introspective in the sense of being 
such that each person must engage in it for himself alone. 
He can not show his results to anybody else. A good bit ot 
ingenuity may be exercised on it and the outcome will be 
found to be rather surprising if not very conclusive as to the 
purpose for which the experiment- was tried. But the results 
as reported may be of some value. Personally, I believe I 
can follow my consciousness down to where I can recognize 
its most basal remaining "content" to be an awareness of 
what I may call extension without definite limitations. It 
seems to me I can detect something to which I could not, 
from its nature alone, apply the terms "I" or "me" as some- 
thing differentiated from everything else. Possibly what I 
note is wholly fanciful, but I seem to feel myself in about 
the condition of psychical life which I imagine a star fish is in. 
Of course I realize how far such a statement is from being 
purified of all thought and other ordinary mental elements. 
Nevertheless, I believe it to be of some value as evidence 
that consciousness is an attribute of the organism as a 
whole, and can neither be held to contain an element which 
can exist separately from the organism, nor be restricted 
to any particular part of the organism as the brain or the 
nervous system. There seems to be some evidence "directly 
felt by us ourselves," and that evidence points to this con- 
clusion as to the nature and "seat" of consciousness. The 
point is susceptible, I am quite sure, of rather rigid experi- 
mental examination. However, the further experiments 
which have suggested themselves to me involve difficulties 
more formidable than I have thus far been in position to 


The reader acquainted with James's notable Chapter X, 
"The Consciousness of Self" (The Principles of Psychology, 

310 The Unity of the Organism 

Vol. 1) will recognize the difference between such introspec- 
tive experimentation as that here indicated, and that so il- 
luminatingly described by James as tried on himself. While 
James's undertaking was to give an account of the thought 
and other processes in consciousness as he could observe them 
in himself, what I want to accomplish requires me to get rid 
of, to ignore as far as possible, the very things which James 
was studying. I want to find whether any "content of con- 
sciousness" remains after thought and the other usual men- 
tal contents are out of the reckoning. I believe, however, 
that James opens the way to such an hypothesis as mine. 
Thus in a footnote we read, "The sense of my bodily exist- 
ence, however obscurely recognized as such, may then be the 
absolute original of my conscious selfhood, the fundamental 
perception that I am. All appropriations may be made to 
it by a Thought not at the moment immediately cognized by 
itself. Whether these are not only logical possibilities but 
actual facts is something not yet dogmatically decided in 
the text." 6 

Except for a little misgiving arising from uncertainty as 
to the exact meaning of "Thought" in this quotation, I be- 
lieve my hypothesis does what James says his text leaves un- 

This foot-note of James's may serve as a switch key to 
shift the current of our discussion from the psycho-con- 
scious phase of life through the psycho-physical to the purely 
physico-chemical phase. The course along which this shifting 
will run can be designated thus: full-fledged intellect (al- 
ready examined), instinct, emotion, bio-physico-chemical or- 

Instinct and Physical Organization 

The discussion from which we have just turned of the 
relation between "inner" and "outer," between "subjective" 

Sketch of an Organismal Theory of Consciousness 311 

and "objective," must be regarded as meeting the require- 
ments of this sketch so far as the first member of the series 
is concerned ; and the relation between instinct and physical 
organization will now receive attention. The evidence of 
vital connection here is so abundant and clear-cut, and the 
views of competent observers are so unanimous that the sub- 
ject can be disposed of quite summarily. Probably the most 
indubitable single block of evidence comes from nest-building 
and cocoon-spinning insects. Many of the facts from this 
field have been so much exploited for the very purposes to 
which w r e now invoke them that a few quotations from and 
remarks upon the writings of naturalists generally acknowl- 
edged for learning and judicious thinking will suffice. 

We turn first to W. M. Wheeler, and take to begin with, 
words which he in turn quotes from Bergson : "As Bergson 
says," we read, " 'It has often been remarked that most in- 
stincts are the prolongation, or better, the achievement, of 
the work of organization itself. Where does the activity of 
instinct begin? Where does that of nature end? It is im- 
possible to say. In the metamorphoses of the larva into the 
nymph and into the perfect insect, metamorphoses which 
often require appropriate adaptations and a kind of initia- 
tive on the part of the larva, there is no sharp line of de- 
marcation between the instinct of the animal and the organiz- 
ing work of the living matter. It is immaterial whether we 
say that instinct organizes the instruments which it is going 
to use, or that the organization prolongs itself into the in- 
stinct by which it is to be used.' And Wheeler continues : 
"The spinning of the cocoon by the larval ant is a good 
example of the kind of instinct to which Bergson refers. 
From one point of view this is' merely an act of development, 
and the cocoon, or result of the secretive activity of the seric- 
teries and of the spinning movements of the larva, is a pro- 
tective envelope. But an envelope with the same protective 
function may be produced by other insect larvae simply as a 

The Unity of the Organism 

thick, chitinous secretion from the whole outer surface of the 
hypodermis. Here, too, we have an activity which, though 
manifested in a very different way, is even more clearly one 
of growth and development. And when the workers of 
(Ecophylla or Polyrliachis use their larvae for weaving the 
silken envelope of the nest, as described in Chapter XIII, 
we have a further extension and modification of the cocoon- 
spinning activities. In this case the spinning powers of the 
larva are utilized for the purpose of producing an envelope, 
not for its individual self, but for the whole colony. In 
conventional works this latter activity would be assigned a 
prominent place as a typical instinct, the spinning of the 
cocoon might also be included under this head, but the form- 
ation of the puparium, or pupal skin, would be excluded 
as a purely physiological or developmental process, yet this 
last, no less than the two other cases, has all the fundamental 
characteristics of an instinct." 

Then immediately follows this statement, especially signi- 
ficant for the proposition of our hypothesis which assigns to 
the individual organism the chemical value of an elementary 
substance: "Viewed in this light there is nothing surprising 
about the complexity and relative fixity of an instinct, for it 
is inseparably correlated with the structural organization, 
and in this we have long been familiar, both with the de- 
pendence of the complexity and fixity of parts on heredity 
and the modifiability of these parts during the life-cycle 
of the individual. Fixed or instinctive behavior has its 
counterpart in inherited morphological structure as does 
modifiable, or plastic, behavior in well-known ontogenetic 
and functional changes." 

The statement that surprise is largely taken away from 
such elaborate manifestations of instinct as those here de- 
picted, by recognizing that the instincts are "inseparably 
correlated with structural organization" and have their 
"counterpart in inherited morphological structure," will, no 

Sketch of an Organismal Theory of Consciousness 313 

doubt, receive the assent of most zoologists, as will also the 
statement that our long familiarity with structural organi- 
zation and morphological inheritance is what makes us re- 
gard these without surprise, and, by inference, as compre- 
hensible. It is not that the corporeal form and structure of 
the worker ants and of the larva? which they manipulate as 
spinning instruments and shuttles for making the nest, are 
necessarily simpler and, on that account, more comprehen- 
sible than are the instinctive acts of the workers, but that 
during our whole lives we have been familiar with structure, 
and ourselves exist as "structural organizations." This is 
equivalent to saying that we have always been not only learn- 
ing but directly experiencing interdependences and correla- 
tions among the common body-parts arid body-acts, and so 
regard them as comprehensible, as explicable. To compre- 
hend really an external complex of structures and activities 
is to live the counterpart of it. To understand such a com- 
plex scientifically is to understand it through a course of 
observation and reasoning; that is, rationally. To explain 
such a complex is to bring in, or recognize consciously one by 
one the constituent elements of the complex, and recognize 
all these as parts of the ensemble. It is to recognize the 
elements in both their isolate and integrate capacities. 

So much for the evidence of integration between instinct 
and physical organization as presented by one carefully phil- 
osophical naturalist. Several other naturalists have gone 
nearly as far, but this single instance is so typical and conclu- 
sive as to the objective facts that it will suffice. In com- 
menting on the significance of being surprised at such rarely 
witnessed performances as those furnished by these ants, 
while we are not surprised at common structures and acts 
of equal or greater complexity furnished by more familiar 
animals and by ourselves, I go beyond, though only a little 
beyond Wheeler. 

The only other zoologist to whom I turn for evidence of 

The Unity of the Organism 

vital relation between instinct and structure is C. O. Whit- 
man. His testimony supplements Wheeler's in that it is 
more exclusively and radically objective than is Wheeler's; 
that is, it verges less toward the subjective- type of presenta- 
tion and draws nearer to the bio-chemical ground work. Al- 
though Whitman wrote relatively little on animal behavior, 
that little seems to me to contain some of the most important 
observations and conclusions which have been produced in 
this branch of zoology. What I utilize is taken from his 
address Animal Behavior. The animals upon which Whit- 
man's chief studies were made were leeches of the genus Clep- 
sine; a salamander (Necturus) ; and pigeons of several spe- 
cies. Our purpose will be best served by quoting a few sen- 
tences which go direct to the heart of the question in hand, 
that namely of the^ vital connection of instinct and basal 
physical structure. "The view here taken," Whitman writes, 
"places the primary roots of instinct in the constitutional ac- 
tivities of protoplasm and regards instinct in every stage of 
its evolution as action depending essentially upon organiza- 
tion". 8 Then, apparently to clarify and emphasize the last 
clause about the dependence of instinct or organization, he 
adds a footnote thus : "Professor Loeb refers instinct back 
to '(1) polar differences in the chemical constitution in the 
egg substance, and (2) the presence of such substances in the 
egg as determine heliotropic, chemotropic, stereotropic, and 
similar phenomena of irritability.' According to this view, 
the power to respond to stimuli lies in unorganized chemical 
substances, and the same powers exist in the adult as in the 
egg, because the same chemical substances are present. Or- 
ganization serves at all stages merely as a mechanical means 
of giving definite directions to responses. 

"The view I have taken regards instinctive action as 
organic action, whatever be the stage of manifestation. The 
egg differs from the adult in having an organization of a 
very simple primary order, and correspondingly simple pow- 

Sketch of an Organismal Theory of Consciousness 315 

ers of response. Instinct and organization are, to me, two 
aspects of one and the same thing, hence both have onto- 
genetic and phjlogenetic development." 

These statements show, as do those given in our discussion 
of the cell-theor}*, how far Whitman went away from full- 
fledged elementalism and toward organismalism. But his 
treatment of instinct and animal behavior reveals what his 
treatment of the cell-theory does not, at least so clearly ; 
namely, how far he also went on the way to the natural his- 
tory mode as contrasted with the mechanistic mode of phil- 
osophizing on biological phenomena. And this gives me a 
pleasant opportunity to testify to the genuinely naturalist 
current that ran through his life and work. An unforgettable 
visit which I had with him among his pigeons not long before 
he died, permitted me to see something of the character and 
depth of his interest in those animals. His whole attitude 
toward them his wonderfully broad information about, and 
understanding of their general ways of life and personal 
idiosyncrasies, his solicitude for them, and his measured af- 
fection for them was such as is never displayed by any 
one who has not very much of the real naturalist about him, 
in his personality as well as in his knowledge. The individual 
pigeons, many of them at aiw rate, appeared to be realities 
to him in a deep sense and not merely "mechanical means for 
giving definite directions to responses" of chemical sub- 
stances. But after all this is said, it must also be said that 
there is no evidence that Whitman ever grasped fully the con- 
ception that the "constitutional activities of protoplasm" in 
which he believed instincts to be rooted, must be the consti- 
tutional activities of protoplasms (protoplasm in the plural 
number), because no individual pigeon is either any other in- 
dividual nor even exactly like any other; and also that the 
existence of protoplasms is dependent upon the organisms 
to which they belong as well as upon the chemical substances 
of which they are composed. Whitman went so far on the 

316 The Unity of the Organism 

road toward organismalism as to believe genuinely in the 
organic and organisation, but not far enough to make him 
accept unreservedly individual organisms. 

We are able- to state definitely wherein lies the great and 
rather unique merit of Whitman's investigations on animal 
behavior. (1) By a judicious combination of pure observa- 
tion and observation aided by experiment and conception, he 
pushed psychic phenomena in the form of instinct down al- 
most to the physico-chemical level ; that is, to the proto- 
plasmic level. (2) He at the same time remained positively 
within the organic, the living realm. His merit is that of 
restraint as well as of positive achievement. He did not per- 
mit his enthusiasm for physical explanation to betray 
him into adopting a phraseology which, while sounding like 
an explanation of instinct, amounts in reality to a denial or 
a repudiation of it. 

So much for the evidence of vital connection between in- 
stinct and organization. According to the schedule indi- 
cated a few pages back for reviewing systematically this con- 
nection through the entire range of psychic life, we have 
next to glance at the connection between the emotions and 

Emotion and Physical Organization 

Approaching this subject as we now are from the direction 
of psychology proper, the well-known James-Lange interpre- 
tation of emotion comes immediately to mind. It will be 
advantageous for our sketch not to focus attention too close- 
ly on any theory or discussion but to take in as much as we 
can of the entire field, keeping in the foreground our own 
personal experiences and observations as contrasted with the 
descriptions and views of authorities. What I mean is that 
the reader shall take himself in hand for serious study as to 
his emotional life, watching himself from hour to hour, day 

Sketch of an Organismal Theory of Consciousness 317 

to day, and year to year under all the varied conditions, 
happenings, purposes, and impulses to which he is subject. 
In doing this a special point should be made of looking back 
scrutinizingly at experiences of particular satisfaction, ela- 
tion, joy, sorrow, irritation, anger, fear, dread, humiliation, 
and shame, as soon after their occurrence as possible that 
they may be fresh in memory. But incidents and episodes 
of one's remoter past which stand out with special vividness 
from the intensity of the particular emotions when they were 
experienced, or because of results which flowed from them, 
will be found illuminating. 

To what extent and in what particular fashion was our 
bodily organization implicated in the feelings and emotions 
we experienced, is our problem. Fortunately one can "live 
over again" as we say; can "work himself into" rather pro- 
nounced emotional states, through a combination of memory 
and imagination. That is, he can be much of a genuine dram- 
atist when all alone, as touching events and scenes of his 
own past experience. What happens to your body when you 
do that sort of thing? is the central question before us. The 
very criterion by which you answer this question you will 
find will be that of how far the body-manifestations appro- 
priate to the particular emotions are elicited through your 
efforts. If your hands do not clinch somewhat, if many of 
your arm, leg, and abdominal muscles do not contract some- 
what, if your respiration does not quicken somewhat, and 
other manifestations, various corporeal indices of anger, do 
not appear quite independently of direct intention on your 
part, you will be sure you have not "worked up" a genuine 
state of anger. The only real knowledge of an emotion is a 
lived knowledge of that emotion. In order to be a true actor 
your body parts must act, directly, automatically, spon- 
taneously, so far as any rational purpose is concerned. And 
what is true of anger is clearly true of all other emotions. 
Our emotional activities may be described as instinctive 

318 The Unity of the Organism 

and reflex activities, the feeling-impulse of which comes 
through intelligence, but is not of intelligence is not under 
the direct guidance and control of intelligence. According 
to this interpretation no animal, no matter how highly con- 
stituted as to instincts and reflexes, could have emotion un- 
less it had intelligence. Emotional activity is instinctive and 
reflex activity of an intelligent organism, with, however, the 
element of intellect eliminated or in abeyance for the time 
being as regards these particular acts. This is what I would 
call the natural history description of emotion. And I be- 
lieve it is in essential accord with James's conception of emo- 
tion, but his description is a psycho-physiological rather 
than a natural history description. I am quite sure that 
what I have just said means virtually the same as the follow- 
ing: "// we fancy some strong emotion, and then try to ab- 
stract from our consciousness of it all the feelings of its 
bodily symptoms, we find we have nothing left behind, no 
'mind-stuff' out of which the emotion can be constituted, 
and that a cold and neutral state of intellectual perception 
is all that remains." 9 


I will now point out wherein I believe the natural history 
description and interpretation of emotion are somewhat truer 
and better than those given by James and other physiologi- 
cal psychologists and, I may add very much truer and 
better than those given by certain writers who approach the 
subject from the physiological side pure and simple. James's 
epigrammatic statements about being afraid because we 
tremble when we meet a bear in the woods ; about being sorry 
because we cry ; about being angry because we strike, do his 
own position some injustice, I think. This is an instance in 
which his gift for piquant writing succeeded too well. But 
the fact ought to be noticed that what he actually says is 
that as between the usual statement, namely, that we tremble 
because we are afraid, cry because we are sorry, strike be- 
cause we are angry, and his way of stating the case, his way is 

Sketch of an Organismal Theory of Consciousness 319 

"more rational." It is only relative, not absolute truth, he 
is aiming at in these statements. Nevertheless, after due al- 
lowance is made for an expressional miscue to some extent, 
there is yet substantial defect in his presentation. Speaking 
in general terms, the defectiveness is not so much in the 
antithesis set up as in the restrictedness implied. Or, bring- 
ing the criticism around toward our particular standpoint, 
the statement falls short of being organismal. 

Cannon has, I believe, indicated the direction in which the 
adequate statement lies. He writes : "We do not 'feel sorry 
because we cry,' as James contended, . but we cry because 
when we are sorry or overjoyed or violently angry or full of 
tender affection when any one of these diverse emotional 
states is present there are nervous discharges by sympathe- 
tic channels to various viscera, including the lachrymal 
glands. In terror and rage and intense elation, for example, 
the responses in the viscera seem too uniform to offer a satis- 
factory means of distinguishing states which, in man at least, 
are very different in subjective quality. For this reason I 
am inclined to urge that the visceral changes merely contri- 
bute to an emotional complex more or less indefinite, but 
still pertinent, feelings of disturbance in organs of which 
we are not usually conscious." 10 What Cannon's criticism 
amounts to, expressed in other language is: while freely 
granting that organs and functions in the usual physiologi- 
cal sense play an essential part in emotion, neither the vis- 
ceral nor any other single set of organs is sufficient to account 
for the whole of any emotion. Visceral changes contribute 
to the "emotional complex," but the real source of the feel- 
ings involved is embedded elsewhere and more broadly in the 
organization. Cannon suggests : "the natural response is a 
pattern reaction, like inborn reflexes of low order." 'The 
typical facial and bodily expressions," he writes, "automati- 
cally assumed in different emotions, indicate discharge of pe- 
culiar groupings of neurones in the several effective states." 

320 The Unity of the Organism 

Without stopping to examine this language in detail, our 
aim will be achieved by pointing out that the more closely the 
various emotions are scrutinized, and the more effort there 
is made to refer them to their causes, the more varied are they 
found to be, and the more widely are we led to search in the 
organization for causal factors. The mental attitude of per- 
fect openness toward any and all facts, both of effect and 
cause, which may occur in a given organic situation, is one 
of the leading characterizations of the organismal conception. 
The assertion that the organism as a whole is the causal ex- 
planation of an emotion or an "emotion complex" is justified 
by two considerations: (1) Except for the organism viewed 
alive and whole and under both its ontogenic and phylogenic 
aspects, the emotion would not exist; and (2) so wide-spread 
and subtle does common observation recognize the parts of 
the organism involved to be in many of its emotional activi- 
ties that for practical purposes, it is better to work on the 
hypothesis that all parts of the organism are implicated than 
to adopt the alternative hypothesis that certain parts only 
are involved ; that is, that some parts are not involved. 

As a matter of fact, I believe that in spirit James' hypo- 
thesis is organismal even though, probably from his training 
and career in formal anatomy, physiology, and psychology, 
he never became entirely free from the Body-Soul antithesis 
and the dogmatisms of "nerve physiology," which have so 
dominated modern physiology and psychology. This opinion 
I base on the general tenor of his discussions particularly 
of the emotions, rather than on his direct formulation of his 
theory of emotion. I will quote a few passages that seem 
particularly to trend in this direction. "No reader of the 
last two chapters [Tlie Production of Movement, and In- 
stinct] will be inclined to doubt the fact that objects do 
excite bodily changes by a preorganized mechanism, or the 
farther fact that the changes are so indefinitely numerous 
and subtle that the entire organism may be called a sound- 

Sketch of an Organismal Theory of Consciousness 321 

mg-board, which every change of consciousness, however 
slight, may make reverberate. The various premutations and 
combinations of which these organic activities are susceptible 
make it abstractly possible that no shade of emotion, how- 
ever slight, should be without a bodily reverberation as 
unique, when taken in its totality, as is the mental mood it- 
self. The immense number of parts modified in each emotion 
is what makes it so difficult for us to reproduce in cold blood 
the total and integral expression of any one of them. We 
may catch the trick with the voluntary muscles, but fail with 
the skin, glands, heart, and other viscera." I ask the read- 
er to make special note of the part of the quotation be- 
ginning, "The various permutations" as we shall have more 
to say about it a few pages farther on. 

Again we read : "Our whole cubic capacity is sensibly 
alive ; and each morsel of it contributes its pulsations of feel- 
ing, dim or sharp, pleasant, painful, or dubious, to that sense 
of personality that every one of us unfamiliarly carries with 
him. It is surprising what little items give accent to these 
complexes of sensibility." 13 I hope the reader will notice 
how easy it would be for me to contend that these state- 
ments come near to my statement about "inner" and "outer," 
or subjective and objective; and also to my formal hypo- 
thesis as to the nature of consciousness. However, I do not 
wish to make too much of such a contention, though I shall 
bring up the point again presently. All I want to do just 
here is to make still clearer the meaning of my view that 
James was organismal in spirit, though not wholly so in for- 
mal statement. To me one of the strongest evidences of 
this was his obvious effort, as indicated by these and many 
other passages in many other writings than his Psychology, 
to describe fully the phenomena with which he chanced to 
deal. As I have remarked in substance so many times in this 
book, one of the most unmistakable signs of the elementalist 
attitude in biology is incomplete and more or less perverted 

The Unity of the Organism 

description. And nowhere, perhaps, in the whole biological 
realm is there a better chance for description of the genuine- 
ly natural history, organismal kind the kind a cardinal 
motto of which is "neglect nothing," than in this very field 
of human emotions, especially of one's own emotions. Nor 
can I refrain from reminding the reader that one of the 
master works in this field is Darwin's The Expression of the 
Emotions in Man and Animals, 27 and that while a leading 
motive of its author was to interpret the emotions in ac- 
cordance with the theory of descent and the natural selec- 
tion hypothesis, probably the most lasting value of the work 
is from its fullness and excellence as a natural history de- 
scription of the emotions and their objective expression. 

As to the fact of vital interdependence between psychic 
life and physical life through the emotions, personal experi- 
ence and observation, backed up and supplemented by many 
authoritative writings, among which those of Darwin and 
James stand out strongly, there seems no longer any room 
for question. The role of the emotions as between "Body" 
and "Soul" may be crudely likened to the splice which a skill- 
ful sailor weaves into two pieces of rope in joining them so 
that there shall be no knot and as great strength as in any 
other part of the rope. In the recent period of psychology 
of so-called physiological psychology we have frequently 
heard about psychology "without a Soul;" and such an idea 
has seemed repugnant to many persons. But if we could 
show that this modern psychology is "without a Body" by 
the same token that it is "without a Soul," the legitimate mis- 
givings about the soullessness of the psychology ought to be 
allayed. And really the organismal conception of psychic 
life is seen, especially when we examine it in the phase of the 
emotions, to amount to such a composition of the Body-Soul 
antithesis. "Body" we can see, as it figured in the old psy- 
chology, virtually signified what we usually mean by corpse, 
or cadaver. "The Body," in that sense was not alive at all. 

Sketch of an Organismal Theory of Consciousness 323 

It was not alive because all the life was taken out of it (by 
the theoretical antithesis) and put into "The Soul." 

Glance at the Equilibrative Interaction Between "Body" and 

"Soul 9 ' 

Going forward from such predominantly observational 
descriptions of psychic life in its emotional phase as those of 
Darwin and James, to such experimental descriptions as 
those being produced by the investigations of Pawlow, of 
Crile, and especially of Cannon, we are getting considerable 
insight into the rationale of how "Body" and "Soul" vitalize 
each other. Modern researches on the physiology or the 
psychology (which one calls it depends entirely on the direc- 
tion of his approach) of psychic life is revealing something 
of the why and how of the poet's instinctive perception, "Soul 
needs Body as much as Body needs Soul." Only one aspect 
of this "why and how" need be noticed in the present discus- 
sion. That is the fact of the balancing off of antagonistic 
emotions to make the normal emotional life just as reflex- 
actions and instinctive actions are largely phenomena of 
equilibration, or balancing-off. 

It should be recalled that we have found this antagonistic- 
equilibrative principle to run through the entire neuro-psy- 
chic life. In the strictly reflex phase the mode of operation 
of the opposing muscles, the flexors and extensors of the 
limbs, as brought out by Sherrington, was cited as a good 
illustration of the principle. A manifestation of the prin- 
ciple in a broader way, as measured by the extent of organic 
parts involved, was seen in the relation of the vagal (cranial) 
and splanchnic (thoracico-lumbar) autonomies, as empha- 
sized by Cannon (Chap. 19, this book) this illustration being 
chiefly in the reflex phase. In a yet higher phase we saw, 
again from Cannon's work, the principle in operation 
through the emotions (Chap. 23) thus bringing it up to the 

324 The Unity of the Organism 

phase of lower conscious life. 

The reader should not forget the insistence throughout 
our presentation of these antagonistic phenomena, that al- 
ways the oppositions and antagonisms and competitions are 
fundamentally constitutive as to the normal organism. Even 
the most pronounced of them are yet in the interest of the 
organism as a whole. They are always partial phenomena 
relative to the whole organism. They have evolved in strict 
accordance with and sub-ordination to the fundamental na- 
ture of the organism in its totality. The opposing muscles 
of our limbs can not break or tear one another under normal 
conditions. Even antagonisms among the parts of the or- 
ganism are possible because the parts belong to the organism. 
The antagonisms of the parts do not produce the organism, 
primarily, but are themselves produced by the organism, or 
at least, are a portion of the means or methods by which the 
organism lives and enlarges, develops and functions. All this, 
be it noticed, holds not merely as touching purely physical 
organization * but as to the entire gamut of psychic life, 
at least up to and including instinctive and emotional life. 

Support of the Hypothesis by the Physico-Chemical Con- 
ception of the Organism 

This prepares us for the final step of switching the discus- 
sion from the psycho-conscious aspect of life to the bio- 
physico-chemical aspect. The place in our discussion to 
which this return naturally takes us is that wherein we con- 
sidered the organism's chemical nature as interpreted by phy- 
sical chemistry. That interpretation has been presented by 
several physiologists but with special insight and cogency by 
F. G. Hopkins. For example, our citation in Chapter 4 of 
the statement that the conception of the organism as a 

* Recall the discussions of growth and chemico-functional integration, 
chapters 17, 18, and 19. 

Sketch of an Organismal Theory of Consciousness 325 

chemical laboratory "is rapidly gaining ground," should be 
recalled, as should also the opinion of Hopkins : "the chem- 
ical response of the tissues to the chemical stimulus of foreign 
substances of simple constitution is of profound biological 
significance," and that further study of the phenomena "must 
throw vivid light on the potentialities of the tissue labora- 
tories." So far this chemical laboratory conception of the 
tissues may be said to be strictly chemical; but let us recall 
what the interpretation is when it passes from chemistry in 
the exclusive sense to physical chemistry and becomes more 
specific as to the laboratory apparatus, as one may say, 
through which the "tissues" work. In other words, recall 
the conception of the cell and its mode of operating, as 
viewed by physical chemistry. The quotations given in 
Chapter 4 may well be repeated in part : ". . . the living 
cell as we now know it is not a mass of matter composed of a 
congregation of like molecules, but a highly differentiated 
system ; the cell in the modern phraseology of physical chem- 
istry, is a system of coexisting phases of different consti- 
tutions." 15 Then from this review our own contention, set 
forth especially in Chapter 7, that wherever in such state- 
ments as those just quoted from Hopkins "the term cell oc- 
curs the term organism really ought to be used." 

It is important for our cause generally that the full 
weight of our argument in support of the view that on the 
strictly physical plane, the organism rather than the ceU 
is really the equilibration system toward which physico- 
chemical knowledge is tending, should be in the reader's con- 
sciousness. At this point if, consequently, this is not so, 
he is urged to read what is said on the point in Chapters 
4 and 7 especially. 

Our central purpose now is to show that the organismal 
hypothesis of consciousness articulates directly and natur- 
ally with the same conception of the organism. Undoubtedly 
it is in the emotional phase of psychic life that this articu- 

The Unity of the Organism 

lation is most open to common observation. Compare, for 
example, James' "Our whole cubic capacity is sensibly alive ; 
and each morsel of it contributes its pulsations of feeling, 
dim or sharp, pleasant, painful, or dubious, to that sense of 
personality that every one of us unfamiliarly carries with 
him," with Hopkins' "On ultimate analysis we can scarcely 
speak at all of living matter in the cell ; at any rate, we 
cannot, without gross misuse of terms, speak of the cell- 
life as being associated with any one particular type of mole- 
cule. Its life is the expression of a particular dynamic equil- 
ibrium which obtains in a polyphasic system . . . 'life' as we 
instinctively define it, is a property of the cell as a whole, 
because it depends upon the organization of processes, upon 
the equilibrium displayed by the totality of the coexisting 
phases." Also compare Hopkins' statement that among 
the different "phases" of the cell in which its life inheres, 
"are to be reckoned not only the differentiated parts of the 
bio-plasm strictly defined (if we can define it strictly), the 
macro-and-micro-nuclei, nerve fibers, muscle fibers, etc., but 
the materials which support the cell structure, and which 
have been termed metaplastic constituents of the cell," with 
James' "each .morsel" of our cubic capacity "contributes its 
pulsations of feeling, etc." 

The congruity of these statements is apparent even when 
taken as here exhibited ; that is, each as standing by itself 
at about the two extremes of the scale of life. When, how- 
ever, they are viewed in connection with my general argument 
that "cell" in Hopkins' statement ought to be replaced by 
"organism" ; and in connection with what we have learned 
from Cannon and others about the mechanism by means of 
which the organism operates in the phase of conscious emo- 
tion, it seems as though our organismal hypothesis of con- 
sciousness comes near to a demonstration. And so far as 
ordinary descriptive natural history is concerned, I believe 
this to be true. However, I recognize, keenly enough, that 

Sketch of an Organismal Theory of Consciousness 

from the standpoint of bio-chemistry, and physiology, and 
also from that of philosophy in the traditional sense, that 
demonstration is not only far away, but is attainable, if at 
all, only by surmounting very formidable difficulties. So I 
reassure the dubious reader that all I am claiming is that 
my two propositions about the nature of consciousness to- 
gether constitute a legitimate scientific hypothesis. 

Personality and Elementary Chemical Substances 

With both the physico-chemical aspect and the psychical 
aspect of our hypothesis now before us more fully and 
sharply than they have been hitherto we will examine an ob- 
jection to it w r hich I apprehend will be the most serious the 
hypothesis will meet ; namely that to the proposition that 
each individual organism has the value in a chemical sense 
of an elementary substance. And since this objection will 
probably be more intolerant and stubborn from the side of 
physics and chemistry than from that of natural history and 
psychology I will adjust my remarks with reference to the 
opposition as thus anticipated. 

The considerations I am going to present might have been, 
in strict expository coherence, presented as a part of my 
discussion of the uniqueness of the individual consciousness 
as marked by its necessary privacy and its difference from 
all other individual consciousness. What we are now to 
emphasize is the fundament ality of objective as contrasted 
with subjective personality of such highly developed animals 
as song birds, domesticable animals, and civilized man. 

A complete definition of "personality" is not obligatory 
for our purpose. Only this much need be said about the 
meaning we shall give the word : First, we deny the right 
claimed by some authors to make personality purely psy- 
chical, or spiritual a thing of the "inner," or "deeper" 
self; "Self" that is, in a thorough-going subjectivistic sense. 

328 The Unity of the Organism 

It is on this ground, as I understand, that some psycholo- 
gists, as G. F. Stout, and apparently C. Lloyd Morgan, 29 
deny personality to animals. All I will say on this question 
here is that I am quite sure that every close observer of the 
higher animals will recognize that if he undertakes to give a 
truly full report of his observations on their behavior he will 
have to speak of the personality of some at least of them 
just as he would of the personality of observed human beings, 
or he will be obliged to call the same thing by some other 
name a kind of procedure against which we have spoken 
strongly throughout this volume. For us, whatever person- 
ality may be, we must conceive it to be founded upon, and 
conformable to, the organism. "Organism" must be the more 
inclusive term. "Person" must stand to "Organism" in the 
logical relation of species to genus. 

Another meaning of personality in this particular dis- 
cussion will concern the uniqueness of each organism as to its 
psychical attributes regarded in their totality. By unique- 
ness I mean not merely the fact that each organism is itself, 
perceptually regarded, but that it is not a replica, a dupli- 
cate of any other. It is not only another organism but it is 
in some measure a different other organism. For the benefit 
of those physical- and metaphysical-minded readers who have 
never informed themselves much about the facts of natural 
history and have never tried seriously to think in the nat- 
ural history manner I would remark that what I have just 
said concerning the uniqueness of the individual organism 
is only re-asserting in a more refined way what botany and 
zoology have recognized more or less definitely since Dar- 
win's time at least, and have partially expressed in the terms 
"individual difference" and "individual variation." 

With this we come to the cardinal point: // individual 
animal organisms, especially individual humans under civi- 
lization, be contemplated with due heed to tlie motto "neglect 
nothing" the conviction will be reached that each and every 

Sketch of an Organismal Theory of Consciousness 

one has literally as much of uniqueness about it as lias an 
elementary chemical substance. 

In order to bring out the truth of this statement we must 
exhibit, in the regular natural history manner, the resem- 
blances and differences between chemical elements on the one 
hand arid the resemblances and differences between human 
beings on the other, and then pool the results of these com- 

To the carrying out of this enterprise the so-called peri- 
odic law in chemistry is of very great importance. The 
essence of this law, stated from the natural history stand- 
point, is that the chemical elements range themselves into 
natural species and genera after much the fashion that plants 
and animals do ; and that the classification is based mostly on 
the chemical attributes of the substances, but partly on their 
physical attributes also. Thus the "halogen group," that to 
which lithium, sodium, and potassium belong, is a genus in 
the sense of descriptive natural history, its species being the 
substances mentioned with others not enumerated. Also the 
group often spoken of in chemical laboratories as "the iron 
group" -the genus containing the species iron, cobalt, nickel, 
platinum, etc., illustrates the point. Two species of the last 
genus, iron and nickel, will be used in our study. Let us 
compare some household utensil made of iron with a similar 
one made of nickel. For the ordinary uses to which these 
implements would be put the difference between the sub- 
stances of which they are made would hardly be noticed. 
The higher specific gravity of nickel (8.5 plus) is so slight 
as compared with that of iron (7.8) that the greater weight 
of the nickel implement would probably not be noticed. Nor 
would the slightly lower melting point of nickel nor its much 
lower magnetic capacity be recognized. The most avail- 
able distinguishing difference is in color, the ordinary house- 
keeper answering you, if you ask how she knows a nickel 
from an iron implement, that the nickel piece is silvery bright 

330 The Unity of the Organism 

while the iron piece is black. 

See now what this means. Actually, as is well known to 
every beginning student in analytical chemistry, these two 
metals are very similar in color as well as in other physical 
attributes so much so, in fact, that some authors apply 
the same term "silver white" to both. What a housekeeper 
really means when she says she knows one implement to be 
of nickel because it is bright and the other to be of iron 
because it is black, is that she is depending on a chemical 
rather than a physical attribute for a distinguishing mark; 
the attribute, that is, in virtue of which iron is acted upon 
much more readily by oxygen in the presence of moisture 
than is nickel. The much greater liability of iron than 
nickel to tarnish and rust is a chemical rather than a phy- 
sical difference between them. This fact, namely that of 
the dependence of distinguishing differences between sub- 
stances more upon chemical than upon physical attributes is 
of very wide applicability in nature, and is greatly impor- 
tant both scientifically and philosophically. 

Now turn from comparing these two elementary chemical 
substances to a comparison of any two human organisms, or 
persons who might be members of a household to which 
the implements might belong. And make the comparison first 
on the basis of the physical attributes just as we began 
comparing the implements of nickel and iron. Does any 
reader doubt that he would find it much easier to distinguish 
the persons than the metals? As to purely morphological, 
that is, physical differences between almost any two persons 
(with the possible exception of certain rare instances of 
"identical" twins), there is no room for question. General 
shape of head, face and features, and the size and propor- 
tions of the various parts of the body furnish many unmis- 
takable distinguishing attributes. 

Sketch of an Organismal Theory of Consciousness 381 

On the Psychology of Subjective and Objective Personality 

But unerring as are the differentiating marks on the 
physical side, such marks are few as compared with those 
on the psychical side. Noting first certain merely physico- 
psychical differences think of the manners of speech and of 
hand writing, to mention only two items ! Undoubtedly these 
differences are to a considerable extent physical but no one 
would seriously question that psychical factors come in all 
along the line. This is perhaps most obvious in speech as 
evidenced by voice modulations, intonations, gesticulations, 
and facial and bodily expressions. Again, differentials are 
everywhere recognizable in responses to sensory stimuli, 
especially in the matter of reaction-time. There are the 
quick and accurate persons, and the quick and inaccurate 
ones ; and there are the slow and accurate and the slow and 
inaccurate types, to go only a step in description and classi- 
fication on this basis. 

Then we proceed to compare the unequivocal psychical 
phases of life: the feeling, the emotional, the esthetic, the 
religious, and the intellectual phases. Here we pass into 
a realm of what might properly be called objective privacy 
in psychology, individuals for the study of which would be 
largely the student's most intimate and most enduring friends 
and associates, human and animal. Such a psychology would 
be undeniably so particular and intimate that much of it 
would be unpublishable even if it had an interest beyond 
the few persons concerned. At the same time there are some 
portions of it of great public importance, one such por- 
tion being exactly what we are in need of in the present dis- 
cussion. I refer to the exceedingly familiar but scientifically 
much neglected definite and sustained psychical differences 
of individuals who by reason of being members of the same 
household or same small community are subject to nearly 
identical influence so far as concerns such fundamental en- 

The Unity of the Organism 

vironic factors as food in the narrow sense, drink, air, light 
and temperature. The duty before us is that of testifying to, 
of msemg, the objectively psychical individual as we did the 
subjectively psychical individual earlier in this sketch. 
'What is needed," writes Sellars, "is not vague statements to 
the effect that individuals cannot be separated or that they 
are aspects of one another, but definitions and analyses." 1T 
Sellars is here raising his voice against the tendency in 
present-day social psychology to make the individual a kind 
of incident in the social order, a by-product of Society. It 
is a satisfaction that the regular course of my psychological 
argument has brought me to where I also may contribute 
something to the definition and analyses essential to check- 
ing the tendency indicated by Sellars. If it can be shown 
biologically and psychologically all in one that personality 
is indubitably objective, both substantively and kinetically, 
not only the Individual but Society will be the gainer, I am 
very sure. For my contribution we will examine in outline 
what may appropriately be called the action-system (adopt- 
ing and expanding Jennings' term) as it manifests itself in 
a small homogeneous group of human beings. Our study will 
be, in other words, one in domestic and neighborhood psy- 

The "material" in this instance must be my own household 
and the handful of persons constituting the colony of the 
Scripps Institution for Biological Research. This group 
is rather specially favorable for such a study in that its 
geographic severance from other groups, and its strictly 
rural habitat give it an exceptionally natural, simple, and 
uniform environment. The analysis might run along any 
one or all of several axes; but our purpose will be accom- 
plished by following one only. That one shall be the reac- 
tion, the behavior, of individual members of the group in 
response to the stimulus of the world war. Were complete- 
ness to be aimed at in the analysis, every individual in the 

Sketch of an Organismal Theory of Consciousness 333 

group would have to be considered. Such a treatment would 
be highly instructive but space limitations forbid us going to 
such length. We must restrict ourselves to a few of the 
more pronouncedly individualistic behaviors and must treat 
even these in a very sketchy fashion. To be remarked at 
the outset is the fact that every member of the group is 
deeply loyal to America and to the cause of the Allies. On 
the very door-sill of the examination we recognize two well- 
differentiated aspects to each person' s action-system, namely 
an aspect of commonality for nearly all members of the 
group ; and an aspect of very pronounced differentiality for 
many of them. 

Behaviors-in-common will receive attention first. In the 
uniform growth, from the very beginning of the struggle in 
August, 1914, of belief in the general rightness of the cause 
of the Entente ; of realization of the meaning of the struggle ; 
and of sentiments and resolutions of devotion to the foreign 
nations with which our nation is finally joined, these experi- 
ences have been very much at one. To be sure this common- 
ness has fallen far short of identity. But as to essentials 
resemblance has been far greater than difference. For ex- 
ample every adult has accepted unhesitatingly his and her 
obligations to the Red Cross ; to the appeals for aid from 
Belgium, France, and the other despoiled countries ; to the 
increasing cost of living; to the buying of Government 
Bonds ; and to the appeals and regulations of the Food Ad- 
ministration. Naturally there has been difference in the 
particular way and extent of response of each in these mat- 
ters ; but in essence there has been nothing differential. 

We turn now to behavior-not-in-common ; behavior, that 
is, which has differentiated the members personally with 
great sharpness. This examination is much more important 
for the subject in hand. The reference here is to each one's 
"bit" as the common phrase had it when our country was 
first entering the conflict. The "war work" (as the expres- 

The Unity of the Organism 

sion has gradually become with the advance toward the cli- 
max of the gigantic struggle) into which each has gravitated 
has much the appearance of the naturalness and inevitability 
presented by the falling of a stone or the flowing of water. 
The case grows so significant at this point that I must par- 
ticularize somewhat more than I have heretofore. A becomes 
an acknowledged leader in "drives" for Red Cross funds, 
Liberty Bond sales, etc. B becomes a regular consultant 
on the knitting of Red Cross articles. C is a highly skilled 
deviser and maker of dishes from "substitute" foods. D 
is appointed an official of the National Food Administra- 
tion. E becomes an official teacher of girls and women as to 
the peculiar duties and obligations of their sex in war times. 
F concentrates nearly the whole of his physical energy upon 
an elaboration of the view that a victory over Germany and 
her allies cannot be really complete without being spiritual 
as well as material- -that the philosophy or theory of life 
being fought for by Germany must be overthrown as well as 
her armed forces. Of the forty adult members of the group 
fully one-half have been incited in a special degree to some 
activity that has a distinct personal character, some of these, 
as above indicated, being very pronouncedly so. The per- 
sonality of these reactions comes to view most distinctly in 
the fact, absolutely certain to an observer whose acquain- 
tance with the persons has been intimate and has extended 
over some years, that no one of those who has settled into 
one of the special, definite, and important pieces of work 
could wholly replace any of the others in their special tasks. 
Probably each could do something at the "job" of any of 
the others were conditions such as to force him to try; but 
success under such conditions would surely be partial, very 
much so in some of the cases. 

This automatic definition and classification of persons sub- 
ject to a common major stimulus, with nearly the same gen- 
eral environic conditions, and with almost complete freedom 

Sketch of an Organismal Theory of Consciousness 335 

of action so far as concerns the particular stimulus, seems 
to me a phenomenon of very great importance since it de- 
pends upon principles of organic beings, especially upon 
principles of civilized man's "being," which are well-nigh if 
not entirely universal, I am sure. Undoubtedly the phenom- 
enon is often much obscured through counteracting ele- 
ments in the environment, especially in social customs, eco- 
nomic conditions and general education among civilized men. 
But in spite of all these, attentive observation will nearly 
always be able to recognize it. Highly significant is it as 
bearing on this particular aspect of the matter, that the 
niches finally found by most of the persons were obviously 
determined to some extent by long continued previous activi- 
ties and unmistakable natural "gifts." 

Another noteworthy fact is the clear indication of not 
mere acceptance, but positive satisfaction on the part of 
most if not all the persons, once they are "settled" to their 
"jobs," this satisfaction prevailing despite the strenuousness, 
perplexity, and wear-and-tear entailed. During the first 
weeks of America's plunge into the maelstrom the anxious 
psychical casting about in our little group, as throughout 
the whole land, presents to the anthropological biologist as 
he looks back upon it a case of trial and error on a gigantic 
scale, the scene being replete with jumbled elements of noble 
zeal, splendid efficiency, mis-expenditure of strength and 
funds, and ludicrous proposals. But out of this, as out of 
this unprecedented instance of world-wide "struggle for ex- 
istence," there is quite sure to come, indeed is coming, as one 
of its first fruits, personality more real and powerful and 
fuller of grandeur than ever. 

While personalities come forth with special distinctness 
of outline and forcefulness of expression during occasional 
events of vast import to the race like the present war in- 
volving literally the whole civilized portion of the human 
species, yet I would insist that the difference between the 

336 The Unity of the Organism 

manifestations at such times and at ordinary times is al- 
most entirely one of degree, rather than of essential nature. 
The attentive observer will not fail to find personalities as 
here understood always and everywhere, no matter how sim- 
ple and lowly the lives, and monochrome the external condi- 
tions. In little details of intelligent, but still more of reflex, 
instinctive, and emotional life, all of which compounded to- 
gether makes what we often call temperament, the keen and 
sympathetic observer will always see persons in the deep 
sense here indicated. Not the transcendent genuises merely, 
the Aristotles, the Shakespeares, the Napoleons, have the 
right to be called personalities, because of the unique powers 
with which they are endowed ; but each and every one of civi- 
lization's humblest-ranked myriads, and each and every 
nature-tutored denizen of the virgin forest, of the untilled 
plain, and of the unregenerate desert, have the same right- 

Personality and the "Breath of Life" Viewed in the Light 
of Physical Chemistry of the Organism 

Swinging the discussion back now on the physico-chemical 
aspect of the organism, I recall first the truth alluded to a 
little while ago, namely, that it is preeminently the chemical 
rather the physical attributes of elementary inorganic sub- 
. stances which furnish the distinguishing marks of these sub- 
stances. Even in the inorganic world we saw that substances 
are most readily and decisively differentiated from one an- 
other by the transformation-products resulting from the 
reaction of the substances upon one another. "Transforma- 
tion of energy," using a form of expression favored by the 
disembodying tendencies in recent chemical theory, is the 
most distinctive thing about all chemistry, inorganic as well 
as organic. The oxidation and other chemically reactive 
changes and products of nickel and iron, we noticed, are the 

Sketch of an Organismal Theory of Consciousness 337 

most differentiative things about these metals. Let us push 
the application of this criterion of difference a little farther 
in comparing human persons. We give energy-transforma- 
tion and work performed a leading place here also. And 
being naturalistically chemical rather than chemically chem- 
ical we are forced to toucli the "high spots" only at first 
regardless of what may be in between them. We are free to 
seize upon the end or completed products of the reactions 
and transformations. What reaction-products, I ask, of 
nickel and iron towards any other substance or set of condi- 
tions are more unlike than the reaction-products of an effi- 
cient Department-of-Justice official, let us say and an ef- 
ficient food conserving house-keeper, in this time of common 
national danger? Yet these diverse products may come from 
not only the same danger stimulus, but likewise from as nearly 
identical physico-chemical environic stimuli as it is possible 
to secure. Were official and house-keeper to eat of the same 
food, drink of the same fluids, breathe of the same air, and 
be subject to the same temperatures month in and month 
out the difference in product would not be a whit less. 

So stands the case when viewed in its "high places" only. 
But the high places are as real places as any whatever. No 
realities, it matters not how obscure or subtle, pertaining to 
the intermediate places, can make the high places other 
than what they are. Judging human beings by w^hat they do, 
by work done through the transformation of the substances 
and energies which they take from the external world, their 
personalities ve surely not less well-attested than are the 
individualities *J elementary chemical substances.* But it 


will not do to be satisfied with touching the high places in 
this rather jaunty fashion. Some attention must be given to 

* A rather full discussion of the point here touched may be found in 
my essay, The Higher Usefulness of Science, where I raise and try to 
answer the query, '"What is nature because man is a part of it?" Per- 
haps a less ambiguous way of asking the question would be, "What must 
nature be in order that it may produce such an animal as man?" 

338 The Unity of the Organism 

the subtler aspects of the problem. The little we shall do 
in this way may be introduced by the query, what reason 
is there for including in our hypothesis the supposition that 
it is "some substance in the air, almost certainly oxygen," 
with which the organism reacts chemically, to produce con- 
sciousness and all other phenomena of life? Why single out 
this substance from the other elementary substances essential 
to life, as for instance carbon or nitrogen? * My reply be- 
gins by recalling the immemorial recognition of the "breath 
of life" the "life giving air" and so on, of universal experi- 
ence. It is well to recall likewise such semi-philosophic con- 
ceptions as that of the pneuma or "psychical breath of life" 
of later Greco-Roman philosophy. The inextricable en- 
tanglement, historically, of breath and air with spirits is 
also worth remembering, especially the continuance of this 
into the modern period of scientific analysis, unmistakeable 
traces of which are seen in the writings of William Harvey 
and the foremost physiologists of the era to which he be- 
longed. For example, the spiritus nitro-aereus of John 
Mayow which, we now know, was his term for oxygen as 
glimpsed first in the history of science, may be mentioned. 

More important than any of these reminders from the his- 
tory of knowledge is that of the familiar fact that the most 
crucial evidences of truly independent or autonomous life of 
the individual higher animal are respiratory. That the new 
born human babe's first breathing-act is its first genuine in- 
dependent life-act is one of the most commonplace of truths. 
And recall how the "return of life" as we say of the nearly 
drowned person, and of one who has "fainted dead away" 
is marked by the resumption of respiratory activities. Cer- 
tain reflexes, as those from stimulating the eyelids, and pos- 

* The argument in answer to this query should be taken as an exten- 
sion of, and in important respects a replacement of, that contained in 
my essay, Is nature infinite? 3 ' 7 wherein I discuss the specificity of in- 
dividual organisms as indicated by how they use their nutrient sub- 

Sketch of an Organismal Theory of Consciousness 339 

sibly certain heart flutterings, may be more persistent move- 
ments than those connected with breathing. But these are 
less certain signs of individual life. It is only to philosophy 
of the elementalist sort that the mere twitch of a hand or an 
eyelid or a trace of heart action would be a satisfactory 
proof of life. Nor would it be to a philosopher of this school 
should the "living substance" under observation happen to 
pertain to a loved relation or friend. Satisfactory evidence 
of life in this case would come only with the nearly simul- 
taneous return of breathing and consciousness. A right 
interesting section could be written at this point on the 
importance of nutriment in the ordinary sense, and of drink, 
as compared with air at the very beginning and ending stages 
of the individual life. For instance such questions would 
have to be considered as that of the independence of the new 
individual for a while at the outset on food-yolk in many 
animals below the mammals, and on placental connections in 
mammals ; that is on material metabolically elaborated by 
the older or parent individual. But such a discussion not 
being indispensable to this sketch, must be foregone. Enough 
here to emphasize the fact that while it may be entirely jus- 
tifiable to regard oxygen as a food as some good modern 
physiologists do the two important facts should never be lost 
sight of that (1) oxygen (air) is the one and only ever- 
present and never varying constituent of the dietary. In 
other words that it is the one constituent which nature sup- 
plies as by "free grace" to use a good old theological ex- 
pression; and that (2) oxygen is the one and only food that 
needs no digesting and so no digestive organs or tissues set 
apart for its metabolic elaboration.* 

Oxygen is the only food which passes directly as such to 

*Were the view held by some physiologists, that the alveolar epithe- 
lium of the lungs transmits atmospheric oxygen to the blood by an active 
process spoken of as a secreting, this statement would need modifying 
somewhat. However, the view does not seem to be accepted by most 

340 The Unity of the Organism 

every part of the organism. In oxygen the organism finds 
one of its most fundamental food materials for which it does 
not normally have to go in search or to compete with other 
organisms. The familiar fact and its significance appear not 
to have attracted the attention of biologists much. Even 
L. J. Henderson 30 who has written so illuminatingly on 
many aspects of organic adaptiveness says nothing definite 
on this point. These two facts are weighty reasons for my 
proposal to look upon oxygen as one chemically elementary 
substance and the organism as another, the reaction be- 
tween which is basal in the production of consciousness and 
all life phenomena. Consequently the problem of how, ex- 
actly, the organism endowed with full-fledged consciousness 
reacts toward oxygen is certainly one of the most important 
of all problems on the purely physico-chemical side of life. 
And, as said early in this sketch, it is just here that my the- 
ory is most avowedly hypothetical. It would be quite out of 
the question to present in the remaining pages of this book, 
even had I the requisite knowledge for doing so, all that 
might profitably be said on the subject. Consequently only 
two or three of what seem to me the most crucial matters will 
be mentioned. 

In the first place I ask the reader to recall what has been 
said in various of the preceding chapters which have brought 
out the indubitable trend of the interpretation of life phe- 
nomena according to the principles of physical chemistry, 
away from the elementalistic conception of the organism. 
The interpretation of the organic cell as a system of phases 
in dynamic equilibrium, so strongly set forth by Hopkins 
and Bayless will be remembered. And this will call to mind 
the sharp way in which the new conception, with its appeal 
to the role of surface-layers, membranes, and areas of con- 
tact between all sorts of constituent substances, sets itself 
over against such pseudo-objective conceptions as that of 
biogens, not to mention the horde of out and out subjectivis- 

Sketch of an Organismal Theory of Consciousness 341 

tic "elements" of which pangens and determinants have per- 
haps had the greatest vogue. The importance of the anti- 
elementalistic tendency of physical chemistry when it comes 
to be applied to biological problems is greatly enhanced, it 
appears to me, by the circumstance that J. Willard Gibbs, 
who was one of the very first to appreciate in a full scientific 
sense the importance of massive as contrasted with minute- 
particle phenomena in inorganic nature, and so was one of 
the "fathers" of physical-chemistry, made no assumptions 
about the invisible composition of substances in his treat- 
ment of "Heterogeneous Equilibrium" and allied topics. 
"Certainly," writes Gibbs, "one is building on an insecure 
foundation who rests his work on hypotheses concerning the 
constitution of matter." 31 If this is true as touching the 
relatively simple structures and movements in the lifeless 
world how much more obviously true is it as touching the liv- 
ing world, and especially such life phenomena as human con- 

sciousness ! 

So we are able to requisition one of the admittedly most 
important advances of modern times in inorganic science as 
support for the supposition that the air we breathe, and 
presumably its oxygen, contributes in some direct and funda- 
mental way to the production of consciousness even though 
this substance, if its "ultimate nature" is what inorganic 
chemistry and physics have hitherto attributed to it, has lit- 
tle or nothing to suggest that it possesses such a unique 
latent attribute. The reader should not fail to recall here 
Hume's recognition of the "secret powers" of substances. 

But is it not possible that physico-chemical and physi- 
ological knowledge of oxygen and air, the "breath of life," 
do contain somewhat more to justify the supposition than is 
usually recognized? In this connection I relate that one of 
the most mentally adhesive statements I ever heard from a 
bio-chemist, its adhesiveness depending largely on the fact 
that the chemist was one of great experience as a laboratory 

The Unity of the Organism 

investigator, was to the effect that chemical analyses make 
known what they find and absolutely no more. In other 
words such analyses never exclude the possibility of sub- 
stances other than those found. And this chemist asserted 
furthermore that all organic analyses leave residues to some 
extent. No manipulative methods are known, it appears, 
capable of effecting a really complete analysis of any or- 
ganic substance. Whether these restrictions on analyses 
still hold I am not sure, though I have seen or heard nothing 
which leads me to suppose they do not. 

It is this general shadow of manipulative imperfection 
which overhangs all formal physics and chemistry, coupled 
with the advances being made from time to time in our 
knowledge of oxygen and air which has led me to put into my 
hypothesis a shade of doubt as to whether oxygen is the con- 
stituent of the air the reaction of which with the organism 
produces consciousness. The demonstration of helium and 
argon, and probably neon, crypton, and xenon in atmos- 
pheric air, all within a little more than two decades, has 
influenced my thinking in the same direction. Besides, the 
idea, become a commonplace of physics and chemistry in a 
single night, figuratively speaking, that the "atom is as com- 
plex as the solar system" has had its part in shaping my 
conceptions ; as have also such well-credentialed conceptions 
from the inorganic sciences as that "Uranium II" is "a long- 
lived element" which is the "parent of the actinium series of 
elements, but has no genetic connection with the uranium 
series" ; and that "in the lead pleiad there are seven ele- 
ments having quite different atomic weights." 

The extent to which, as exemplified by this case, the inor- 
ganic sciences have found themselves driven into the organic 
realm for terms with which to express their new conceptions 
must impress every thoughtful person. Earlier, what we 
might describe as purely contemporaneous physical dynamics 
had to borrow such terms as energy, power, force, work, 

Sketch of an Organismal Theory of Consciousness 343 

from the nomenclature of living beings. Later, with the per- 
meation of all knowledge by the conception of the natural 
or derivative origin of everything (a genuinely organic con- 
ception, notice), has come even for elementary chemical sub- 
stances, the induction into physics and chemistry of such 
ideas as genetic relations, parenthood, and length of life. 
So my suggestion that the air we breathe must be recognized 
to possess latent attributes which by reacting w T ith the or- 
ganism produce consciousness, falls into a genetic series in 
the history of the interpretation of nature. 

The very important question, as already indicated, of ex- 
actly how atmospheric or molecular oxygen operates in the 
living being generally and the conscious being particularly, 
is largely for the future to answ r er. One should never fail, 
however, to couple this question with the same question as to 
the behavior of oxygen, and for that matter of any other 
chemical substance, in any reaction whatever. Exactly how, 
for example, does oxygen operate with hydrogen to produce 
the attribute of ref rangibility of water ; or with phosphorus 
to produce the peculiar glow which that substance may ex- 
hibit under some conditions? 

Concerning the positive knowledge and the views as to 
details of the action of oxygen in connection with the or- 
ganism, only a little can be said here though that little may 
be very important. Looked at from the standpoint of the 
old, the orthodoxly atomistic chemistry, probably the most 
anomalous tiling about my hypothesis is that the organism 
conceived as equivalent, chemically speaking, to an elemen- 
tary substance, is the unquestioned fact that the organism is 
not only composed of several chemical substances, but that 
one of these is oxygen itself. Stated baldly, the anomaly is 
that two chemical substances are supposed to react upon each 
other, one of which (the organism) is known not only not 
to be simple, but to contain the other substance. But even 
the old chemistry with its "compound radicals," of which 

344 The Unity of the Organism 

cyanogen (CN) 2 is said to have been the first discovered, and 
of which the . unitedly-acting combinations of carbon and 
hydrogen as methyl, CH 3 , affords some slight support for 
our conception so far as the mere matter of chemically uni- 
tary compoundedness is concerned. In so far, however, as 
technical chemistry can be drawn upon for supporting our 
hypothesis, it is the new, or physical chemistry, as has 
been repeatedly stated, that is our main reliance. Unless I 
am greatly deceived, the real inwardness of that great move- 
ment in inorganic science is against the age-old conception of 
the ultimate adequacy of atoms to explain inorganic na- 
ture, almost as positively as the organismal conception is 
against the ultimate adequacy of any constituent element 
whatever, to explain organic nature. The surface energies, 
for example, developed at contact faces and giving rise to 
the phenomena of adsorption * appear to be not a whit less 
real and ultimate energies than are any that can be attrib- 
uted to atoms and molecules taken as such. And, be it no- 
ticed, one of the most distinctive things about these areal and 
massive energies is that they dominate atomic and molecular 
energies to a certain extent. This is just what the now uni- 
versally recognized principle of "mass action" is in so far as 
such action has been studied enough to make possible its for- 
mulation into law ; that is enough to learn how it influences 
velocity and quantity of chemical change. But would any 
careful physicist or chemist pretend to know to a certainty 
that such action is restricted to influence of that sort? Surely 
not. Are we certain for instance that it can not under any 

* Adsorption is the loading- of the surface of a solid body immersed in 
a solution, with the dissolved substance. Thus it is by adsorption that 
charcoal takes the coloring matter out of a colored solution. The action 
results from the facts that there is surface tension at the interfaces be- 
tween the charcoal and the liquid, and that this tension is lessened by 
the presence of the dissolved color-substance in the liquid. The sub- 
stance then moves to the place of lessened tension and concentrates on 
the surface of the solid. 32 The principle has very wide application in 
nature, particularly in organic nature, where colloidal substances and 
water are in contact so extensively. 

Sketch of an Organismal Theory of Consciousness 345 

circumstances influence qualitative as well as quantitative 
change? Surely we are not. This of course is far from 
contending that mass action actually does influence quali- 
ties. My sole point is that so long as there is lack of 
certainty that it does not or may not exert such influence 
any assumption which implies such certainty is unwarranted 
and unscientific. 

Putting together, then, the physically massive concep- 
tions of inorganic chemistry and the organismal conceptions 
of bio-chemistry w r hat seems to follow touching the chemico- 
substantive composition of organisms is that a portion of 
all the substances essential to life, carbon, oxygen and others, 
have been combined from all eternity (whatever be the mean- 
ing of the phrase) in the peculiar way called organic, while 
other portions have remained in the state called inorganic. 
This leads me to remark, quite incidentally so far as this 
discussion is concerned, that according to this view the as- 
sumption would be that organisms have always existed, or at 
least that they have existed as long as "matter" or anything 
else of which we have any information or clear conception, 
has existed. The warrantableness of this assumption I am re- 
lieved from arguing here from having treated the problem 
at some length in another place. (Are we obliged to suppose 
the spontaneous generation of life ever occurred?)^ All 
that need be said now about the outcome of that discussion is 
that the warrantableness lies in the absence of any ground 
for assuming the contrary. I take my position squarely on 
the direct evidence in the case. All the evidence of that sort 
we have and in that discussion I emphasize the fact of its 
vast quantity- -is to the effect that organisms are produced 
by other organisms known as parents and in no other way.* 

* To the stock and rather vapid rejoinder that such a solution of the 
problem of the origin of life is no solution at all, but only a putting off of 
the difficulty, the obvious reply from my standpoint is that I am making 
no pretense of "solving the problem," as "solution" would be meant in 
the anticipated rejoinder. From my standpoint, however, the everlast- 
ingly-from-parents hypothesis would be a solution of the problem if the 
hypothesis were proved true. 

346 The Unity of the Organism 

We can now state briefly as much more of the bio-chemical 
aspect of the problem as seems indispensable to our present 
argument. A few remarks on what the physiology of our day 
often calls tissue respiration will compass what is in mind. 
The key fact in this is of two-fold character: (1) The tissues 
of the organism, not its blood or any other fluids, contain the 
substance which is in the strictest sense living. () This 
substance is called living because chemical changes of a very 
distinctive sort are going on in it. These changes are of a 
fundamentally double nature as regards atmospheric or 
molecular oxygen ; namely, combinative and incorporative 
change, and separative and expulsive change. The last-men- 
tioned, the separative and expulsive change, is known as oxi- 
dation and manifests itself to ordinary experience in the dis- 
charge of oxygen combined with carbon as carbon dioxide, 
and in the setting free of energy in the form of muscular and 
other work, and of heat. The first-mentioned, or incorpora- 
tive change, consists in taking in and storing up oxygen, 
"somehow," as the more carefully worded physiologies put 
it. This statement may be taken as a very brief natural 
history description of the most fundamental steps in what 
formal physiology calls metabolism with its two aspects, the 
constructive, or anabolic, and the destructive, or katabolic. 
Probably no one will question that this conception of the 
foundations of the life process for nearly, if not quite, all 
animal life is that held by the best physiologists since the 
time of C. Bernard at least. No physiologist whom I have 
consulted has stated the nature of the process more definitely 
than has Sir Michael Foster. "The Respiration," he writes, 
"of the muscle then does not consist in throwing into the 
blood oxidizable substances, there to be oxidized into car- 
bonic acid and other matters ; but it does consist in the as- 
sumption and storing up of oxygen somehow or other in its 
substance, in the building up by help of that oxygen of 
explosive decomposable substances, and in the carrying out 

Sketch of an Organismal Theory of Consciousness 347 

of decompositions whereby carbonic acid and other matters 
are discharged first into the substance of the muscle and 
subsequently into the blood/' And he points out in other 
connections that what is true of muscle in this regard is es- 
sentially true of all other tissue systems. In another still 
more recent text book we read : "Nothing definite is known, 
however, as to the nature of the probable combinations 
formed by oxygen with the different materials for building 
up muscles and other tissues, or of the intermediate anabolic 
and katabolic forms through which it passes in combining 
with carbon into carbonic acid." And this author then 
expresses what are, apparently, his own views, by quoting 
from Foster as follows : "The whole mystery of life lies hid- 
den in the story of that progress [that of construction and 
destruction in the tissues] and for the present we must be 
content with simply knowing the beginning and the end." 

The kernel of my suggestion so far as metabolism is con- 
cerned, is that the anabolic, or the assimilative, the truly 
synthetic aspect of the complete operation, is the continual 
renewal, or keeping up of the oxygen constituent of the 
organism which comes to it by heredity, that is which has 
alwavs been in the "line of descent." It is the maintenance 


of what might be spoken of as the original oxygen constitu- 
ent of the organism. There would always then be operating 
in the organism oxygen of two sources, that from the one 
source designated, employing our well-established evolutional 
terminology, phylogenic or hereditary oxygen ; and the other 
ontogenic or individual oxygen. In general the same kind 
of reasoning would hold for the other chemical simples, car- 
bon, nitrogen, and so on; but these are in quite a different 
status from oxygen owing to the fact that they are not 
normally taken by the animal organism in the pure or uncom- 
bined state, but only in some other organic combination, 
as food in the ordinary sense. 

Metabolically expressed, then, we may say in short that 

348 The Unity of the Organism 

the warrantableness for considering the individual organism 
as a chemical element, is the fact that it maintains its identity 
as regards all its elementary constituents except one, oxygen, 
be wrenching these, so to speak, from other organic com- 
pounds (by digesting these) and then by synthesizing the 
elements into its own particular substance. Another way of 
expressing the same conception is to say that the organism 
is an element, chemically speaking, because it reacts directly 
in a chemical sense with another element. 

Did this chapter pretend to be anything more than a 
sketch of a theory of consciousness a considerable discus- 
sion of the "activation" of oxygen would naturally come in 
somewhere, perhaps at this point. The essense of activation 
is the fact that when oxygen passes into the organism by the 
respiratory process it is somehow changed into a condition 
which enables it to oxidize living tissue-substances as it 
can not to any degree, seemingly, when brought into con- 
tact with the same substances outside the organism. This 
discussion would involve the various theories which have 
been put forward to account for this phenomenon, as those 
which make use of the principle of enzymes, of peroxides or 
of some other. All that our aims here require us to notice 
is that nothing conclusive as touching the nature of activa- 
tion would come from the discussion. How unsatisfactory 
a state this whole subject is in may be seen from the follow- 
ing words of a foremost American biochemist : "It has been 
a popular practice to appeal to hypothetical enzymes to 
explain some of the obscure chemical transformations in the 
organism. Thus we have been wandering through the mazes 
of the oxidases, oxygenases, peroxidases, reductases, cata- 
lases and other products of perplexing nomenclature in the 
hope of escaping the uncertainties of intermediary meta- 
bolism." 36 

Sketch of an Organismal Theory of Consciousness 349 

Summed-up Statement of Justification of the Hypothesis 

The final gathering-up-and-putting-together may now be 
made of all that has been said about the physico-chemical 
aspect of the organism on the one hand, and about its psy- 
chical aspect on the other. That is to say, we are now ready 
to epitomize the results of our examination of the ancient 
and honorable but withal unsolved problem of how Body and 
Soul go together. As regards "body" or "the physical" we 
have been led to the physico-chemical conception of the or- 
ganism as a well-nigh inconceivably complex mass of sub- 
stances, mostly in the colloidal state, operating as a system 
of phases in dynamic or constantly changing equilibrium. 
As regards "soul" or "the psychical," we have found also a 
series of phases of activities, namely the phases of intellect 
and reason, those of instinct, those of feeling and emotion, 
those of the will, those of the tropisms and the "simple re- 
flexes," and finally those of simple protoplasmic response. 
According to my hypothesis, the phases of the bio-chemico- 
physical sort and the phases of the psychical sort have com- 
mon ground in the organism as a whole, the phases of in- 
tellect and reason corresponding to the cerebro-spinal nerv- 
ous system ; the phase of instinct corresponding probably to 
the autonomic nervous system ; the phases of feeling and emo- 
tion corresponding mainly to the glandular and visceral sys- 
tems ; those of the will to the body-muscular system ; those 
of the tropisms and simple reflexes to the receptor-conductor- 
effector systems ; and finally those of simple protoplasmic 
response to the fundamental protoplasmic mechanism of 
response, whatever its structure. 

According to the scheme presented in the sketch and 
summed up here, just as physical functioning and physical 
form reach back to the very dawning of animal life, both in 
the individual and in the race or type, so consciousness with 
its nether limits in what, following the terminology of em- 

350 The Unity of the Organism 

bryology (see section on the pro-morphology of the egg- 
cell, Chap. 8), might be called pro-consciousness, is an 
attribute of all animal organisms. As comparative anatomy 
and physiology have made us familiar with the physical as- 
pect of the animal organism existing as the fully realized 
or developed adult at one end of the ontogenic series, and as 
the unrealized adult or germ at the other end of the same 
series, exactly so its psychology gradually familiarizing us 
with the realized, or adult mind at one end of the ontogenic 
series, and as the unrealized or germinal mind at the other 
end of the same series. When we affirm that the completed 
individual organism is latent in the germ, we must under- 
stand that the psychical aspect no less than the physical 
aspect is so latent. With very little doubt, it seems to me, 
the real meaning of the so-called sub-conscious, and of 
psycho-analysis as a method of investigating it, is that the 
ontogenic stages of the psychic life of the human organism 
are being discovered and that a method of investigating these 
stages is being worked out. Freud and his followers have 
been and still are somewhat in the dark, I think, as to just 
what they are doing, albeit their discoveries and methods are 
of the utmost importance. 


1. Wheeler, W. M 519 '21. Ritter, W. E. (1918 2 ) Essay 2 

3. Sedgwick & Taylor 262-3 22. Huxley, T. H 98 

4. Royce, J 2 23. Huxley, T. H 96 

5. Royce, J 3 24. Huxley, T. H 100 

6. James, Wm 1-341 25. Huxley, T. H 94 

7. Wheeler, W. M 521 26. Dewey, John 9 

8. Whitman, C. 310 27. Darwin, Chas 

9. James, Wm 11-451 28. Crile, G. W 

10. Cannon, W. B 280 29. Morgan, C. Lloyd 256 

11 . Cannon, W. B 282 30. Henderson, L. J 

12. James, Wm 11-450 31. Gibbs, J. W Pref. x 

13. James, Wm 11-451 32. Lewis, Wm. C. McC...II, 303 

14. Hopkins, F. G 217 33. Ritter, Wm. E. (1918 2 ) Essay 1 

15. Hopkins, F. G 220 34. Foster, M 469 

16. Hopkins, F. G 220 35. Luciani, L I, 395 

17. Sellars, R. W 75 36. Mendel, L. B 21 

18. Descartes, Ren6..Pt. 1, Sec. 8 37. Ritter, Wm. E. (1918 2 ) 

19. Montague, W. P 120 38. Wendt, G. L 442 

20. Hume, David. . .Sec. IV, Pt. 1 39. Stout, G. F 680 


THE argument in favor of the organismal way of viewing 
living nature has now run what appears to me its natural 
course, to its inevitable end. Yet I cannot bring myself to 
write "Finis" without making a few remarks which though 
connected vitally with the argument, do not seem an essential 
part of it. 

These remarks concern the general effect of the organismal 
standpoint on those who may grasp it firmly and adopt it 
unreservedly. Since, as pointed out in the "Historic Back- 
ground" with which this book opens, the standpoint has been 
recognized by biologists with varying degrees of fullness 
from the time of Aristotle at least, there can be no doubt 
that the human mind is naturally attuned, as one might say, 
to this general type of response to organic phenomena. It 
seems therefore fitting that a presentation like that which I 
have made should be accompanied by a few words on the 
probable influence of a wide prevalence of the organismal 
view. The pertinent question will be asked, how could it 
have come to pass that if the standpoint has been so long 
in the world it should have missed full recognition and have 
failed to exert its due influence? The reply is obvious to an 
attentive reader of this book : At no time until the present 
in the long historical growth of knowledge of the living world 
has information been sufficient to make possible a rounded- 
out statement of the conception. To illustrate, it is only in 
the very last years that enough has been known of the 
physical chemistry of the cell to engender such an interpre- 
tation of this exceedingly important biological entity as that 
which biochemists are just now reaching. Yet this interpre- 


352 Postscript 

tation is indispensable to anything even approaching a full 
development of the organismal view. 

But nothing stands out more boldly from the pages of 
this book than the insufficiency even yet, of actual knowledge 
for making the standpoint complete. If therefore, I append 
to my presentation a brief reference to the larger effect the 
view has had on myself, and* on this basis forecast what the 
effect would be on thinking people generally were they to 
make it their own, such a forecast will surely be in harmony 
with the larger purpose of the book, even though the antici- 
patory remarks have no place in the presentation itself. 

The long and laborious gathering and arranging of facts, 
and weighing of natural evidence and formal arguments 
which has constituted the development of the standpoint in 
my own mind, has compelled me to re-examine and re-assess 
the whole frame and fabric of my spiritual life. Nothing, so 
far as I can tell, has escaped. Not my scientific knowledge 
alone my professional stock-in-trade but all my ideas and 
beliefs touching religion, art, society, politics, industry, per- 
sonal relations, and private living, have come in for their 
share of scrutiny and renovation. 

An exceedingly brief "synoptic" classification and char- 
acterization * of the entire range of these effects can be 
given in the terms of formal science and philosophy. 

As to classification, the effects fall into a two-fold group- 
ing. One of the groups appertains to the great province of 
the nature of knowledge; the other to the equally great 
province of the nature of morals. 

The characterization of effects on the nature of knowl- 
edge which seems to me most inclusive and most practically 
significant, may be stated thus: By the validation of ob- 
jective knowledge, largely through the principle of what I 
have called standardization of reality, but partly through 

* See my essay, The Place of Description, Definition and Classifica- 
tion (Ritter, 1918). 

Postscript 353 

the organismal hypothesis of consciousness, such knowledge 
is elevated to the rank of strict equality with "pure thought," 
often so-called; that is, with subjective, or intuitive knowl- 
edge. In this way mathematico-mechanistic science is de- 
prived of the regal place it has claimed for itself since the 
era of Descartes and Leibnitz, and is brought to the plane 
of absolute equality as to importance and dignity, with 
sense-experiential science. By thus adjusting the claims of 
these two great realisms of science, an attitude toward the 
infinite totality of nature, and a methodology for interpret- 
ing it, which have hitherto borne the stamp of subjection 
and inferiority assume their rightful places in the great 
hierarchy of philosophical science. This leveling-down of 
mathematical mechanics and the deductive method and level- 
ing-up of observational knowledge and the inductive method, 
implies the complete overthrow of psycho-physical dualism 
in psychology, and the rescue of personality from bondage 
to a theoretically infinite monotony of "Matter and Energy." 

The characterization of the effects of the organismal view 
on morals centers around the perception that in the establish- 
ment of human personality the persons are organically in- 
terdependent upon one another; that is, interdependent 
through their "attributes of relation," this resulting in the 
incorporation of men into a pluralistic universe far more 
real and vital than philosophic pluralism has hitherto been 
in position to grasp. Through a type of human conduct 
guided by knowledge of these principles of personality and 
the interdependence of personalities, and through supple- 
menting mathematico-mechanistic methods of study by a 
rigid application of observational and statistical methods, a 
genuine science of morals, both theoretical and practical, 
is made attainable. 

That my enterprise of developing the organismal view is 
only part and parcel of the general current of interpretation 
of living nature which has flowed through the centuries seems 

354 Postscript 

clear even from my meager acquaintance with the history of 
philosophic thought. Thus we read in Windelband (A His- 
tory of Philosophy, Eng. by Tufts,) : "For the decisive fac- 
tor in the philosophical movement of the nineteenth century 
is doubtless the question as to the degree of importance 
which the natural-science conception of phenomena may 
claim for our view of the world and life as a whole." (624). 
Then after speaking of the sharp antithesis between the 
Weltanschauung elaborated by the "Highly strained idealism 
of the German Philosophy" of the early nineteenth century, 
and the "materialistic Weltanschauung" of the later decades 
of the same century, the author writes : "If we are to bring 
out from the philosophical literature of this century and 
emphasize those movements in which the above characteristic 
antithesis has found its most important manifestation, we 
have to do primarily with the question, in what sense the 
psychical life can be subjected to the natural-science mode 
of cognition." (p. 625). 

That Part II of this book of mine, especially Chaps. 20 
to 24, go a long way toward answering the cardinal question 
formulated by Windelband appears to me certain. And, I 
may add, it also seems quite clear to me that the gigantic 
struggle at arms which that philosopher's nation has now 
brought upon the world, is one of the strongest proofs that 
philosophic thought and, following this, social and political 
leadership in Germany have failed miserably to discover the 
Via Media between the Weltanschauung of the "highly 
strained idealism of the German Philosophy" and the mate- 
rialistic Weltanschauung which has finally reached its nat- 
ural climax in militaristic brutism, and is almost certainly 
(Sept., 1918) approaching its overthrow. 

Nothing could more fittingly end this book, devoted as it 
is to demonstrating the operative nature of organic unity 
in one of its great segments, than a reference to the fact 
that the philosophy of life now determining German morals, 

Postscript 355 

and which has drawn its inspiration largely from the hypo- 
thesis of natural selection, has failed- -pathetically beyond 
the power of words to express if done unintentionally ; and 
criminally in equal measure if done intentionally- -to under- 
stand the real meaning of Darwin's teaching as a whole. 

Certain it is that had the German philosophers of Maclit- 
politik recognized the place of unqualified supremacy as- 
cribed by Darwin to the mental and moral endowments of 
man, it would have been impossible for them to make the 
dogma of survival of the fittest serve their ends in any such 
way as they have made it, and done so honestly. Attentive 
reading of Tlie Descent of Man makes it perfectly plain 
that Darwin simply accepted all the higher human attri- 
butes moral, esthetic, and religious, no less than those of 
the intellect as fundamental data in his reasoning about 
man's evolution. His sole effort as touching these was merely 
to see in how far they could be regarded either as helped 
forward in their development by natural selection, or at least 
as not inconsistent with it. Apparently it never even oc- 
curred to him to regard his hypothesis as supreme-over-all, 
so that all attributes whatever, the noblest ones of man with 
the rest, must either be forced into conformity with it, or 
their reality and pow r er virtually denied. "I fully subscribe 
to the judgment," runs the opening sentence of the chapter 
on "The Moral Sense," etc., "of those waiters who maintain 
that, of all the differences between man and the lower ani- 
mals, the moral sense or conscience is by far the most im- 
portant." And, especially significant at this time, Darw r in 
quotes with obvious approval, an apostrophe to Duty by 
Kant, in which this "Wondrous thought'' is represented as 
"holding up its naked law" in the soul, and demanding 
reverence. Darwin's entire discussion in this part of the 
Descent makes it clear that w r hat he had in mind was to 
discover as far as possible the germs of "conscience," of 
"feeling of right and wrong," of an "inward monitor," of 

356 Postscript 

"sympathy," of "parental and filial affection," of "social 
affection," of the "instinct of self-sacrifice" and so on, in the 
lower animals so as to have a starting point for these attri- 
butes as they occur in civilized man. It was not at all his 
purpose to show, as the German perversion of the struggle- 
and-survival hypothesis holds, that the evolution of man 
has consisted largely in a farther differentiation and intensi- 
fication of the dominantly brute attributes, with an infusion 
as a kind of by-product from the struggle for existence, of 
certain "humanistic sentimentalities," which in reality are 
signs of weakness and must be suppressed.* 

And this perversion by German science and philosophy of 
Darwin's teaching is rooted very deep in German culture and 
character. The straightforward, common-sense descriptions 
and inductions of the practical-minded, country-dwelling, 
country-loving, unacademic English naturalist were alto- 
gether too simple and unsophisticated to satisfy a Kultur 
permeated through and through with the "highly strained 
idealism" of Kant, Fichte, Hegel, and Schopenhauer. The 
two worst errors committed by Darwin were his over-em- 
phasis on the natural selection hypothesis, and his pro- 
pounding of the gemmule-pangenesis hypothesis ; and it is 
highly characteristic that it was in just these two "strained" 
speculations that German biology and practical philosophy 
should have taken up Darwinism the most ardently and over- 
worked it the most absurdly and disastrously. 

My examination of the germplasm-determinant theory of 
Weismann in Part I of this book has revealed something of 
the scope and nature which the gemmule fallacy was destined 
to assume when it fell subject to German speculation. The 
more subtle and far-reaching and humanly practical conse- 
quences of the adoption and elaboration of the struggle-and- 


The effort which Dr. George Nasmyth has made in his book Social 
Progress and the Darwinian Theory to set right Darwin's position in 
this matter, ought to bear fruit after a while. 

Postscript 357 

survival hypothesis by German speculation has not yet been 
subjected to thorough-going biological criticism, though sev- 
eral moves in this direction have been made. 

Even the realism of recent German political and economic 
theory and practice is a "highly strained" speculative real- 
ism. This philosophical monstrosity is largely attributable, 
demonstrably so I believe, to a cultural and governmental 
system in which the principle of universal organic personality 
is grossly violated. And what a price in misery and blood 
and treasure the whole world, but old Europe particularly, 
is paying for a consummation which a truer philosophy of 
life would have foreseen and forestalled ! 

Can the leaders of German Kultur be convinced of the 
fundamental fallacy of their theory of human and national 
life, only by discovering that their military establishment, 
built up through many decades of patient, costly organiza- 
tion and discipline, but under guidance of a philosophy of 
mechanism and brutism, is yet incapable of overpowering a 
military establishment, a large portion of which may be im- 
provised in the course of a few months, if such improvision 
be under guidance of a philosophy of personality and hu- 
manism ? 



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ACROGEMALY. A disease charac- 
terized by hypertrophy of the 
terminal parts of the body, as 
of the face and extremities; an 
outgrowth involving bony and 
soft parts. 

ADRENALS. A pair of small glands 
situated in front of the kidneys. 
They are glands of internal se- 
cretion, their secretion exercising 
in particular a regulating effect 
on the nerves of the heart and 
blood vessels. 

ADRENIN. The "active principle" 
in the secretion of the adrenal 

ALCYONARIA, include many corals 
and other coelenterate animals 
with eight mesenteries and eight 

ALVEOLAR. In anatomy, a numer- 
ously-pocketed, or sacculated, 
structure, typified by the ter- 
minal cavities of the lungs, but 
occurring in various tissues; be- 
lieved to constitute also one kind 
of protoplasmic structure. 

AMOZBA. A unicellular animal, a 
genus of rhizopodous Protozoa. 

AMPHIBIAN. An animal living 
both in water and on land. 
Properly a class of vertebrates 
whose young are typically aquat- 
ic and respire by gills; examples, 
frogs, toads and salamanders. 

VMPHioxrs, literally pointed, or 
sharp at both ends. The cur- 
rent name for one of the very 
simplest and lowest vertebrate 
animals occurring in the sand 
and mud of the seashore in 
many parts of the world. 

ANABOLIC, the chemical up-build- 


ing of the living body; construc- 
tive metabolism. 

ANTIBODIES. "The products of a 
reaction of the body towards a 
natural or artificial introduction 
into it of certain foreign sub- 
stances, bacteria and their poi- 
sons, vegetable poisons of other 
kinds, and various albuminoids." 
The name antibodies has refer- 
ence to the antagonism these 
products have for the introduced 

ANTIGENS. Substances the reac- 
tion of which with the living 
body produce antibodies. 

ASCIDIANS. Marine animals hav- 
ing a gelatinous or leathery en- 
velope containing cellulose. In 
the larval stage a notochord or 
forerunner of the vertebral col- 
umn is present. Some free liv- 
ing species retain the notochord 
all through their lives. 

AXON. The long, slender, sparce- 
ly-branched, nbrillar process of 
a ganglion cell; contrasted with 
the shorter, more branched, more 
irregular dendron. 

AXOSTYLE. A slender, flexible rod 
of organic substance forming a 
supporting axis for the body in 
some Protozoa. 

BIOGEN. Literally life producer. 
Imaginary ultimate units of life. 
Such special significance as the 
"biogen theory" has over other 
theories which make imaginary 
vital or physiological units a 
goal of the ultimate explanation 
of life, is found in the fact that 
the biojren theory aims to be 
more definitely chemical than 



the others. The German physi- 
ologist Max Verworm has elab- 
orated this speculation more 
fully than has any one else. 

BIOPHOR. Literally life carrier. 
Biophors, the imaginary ultimate 
vital units of the Weismannian 
system of speculative biology, 
differ from biogens in the fact 
that Weismann, not being a 
chemist or even a physiologist, 
but a zoologist interested in re- 
production and heredity, rather 
than in function generally, did 
not undertake to put his specu- 
lation on a chemical basis. 

BIOPLASM. Formative living mat- 
ter; not differing in any but a 
speculative way from proto- 

BIOTIC. Pertaining to living 

BLASTOGEKESIS. Reproduction by 
budding, as used in this book; 
in general, propagation from an 
undifferentiated germinal mass. 

BI,ASTOMERES. The first segments 
or cells formed by the division 
of the ovum. 

BLASTOZOOIDS. The united individ- 
uals produced by budding, and 
constituting the colony, or cor- 
mus in the compound ascidians. 

BLASTTJLA. The stage of develop- 
ment of the embryo from the 


ovum in many animals, in which 
the organism consists of a hol- 
low sphere the wall of which is 
composed of a single layer of 

BRYOZOA. Literally "moss ani- 
mals," from the resemblance, 
fancied more than real, of some 
of the species to mosses; also 
called Polyzoa. Marine animals 
occurring abundantly on all 
shores. Most of the species 
propagate by budding as well as 
by eggs and sperm, the bud-pro- 
duced individuals remaining at- 
tached to one another to form 
colonies, as in many hydroids and 

ascidians. Each individual con- 
sists of a body proper bearing 
a circle of tentacles, and an en- 
veloping case often calcareous, 
into which the body may be 
quickly and completely re- 

CALYMMA. The much-vacuolated 
portion of the body of radio- 
laria, situated outside the central 
capsule, the vacuoles containing 
fluid impregnated with gas. The 
main office of the structure 
seems to be in connection with 
the flotation of the animals. 

CAMBIUM. The layer in woody 
plants between the outer dead 
layer, or bark, and the inner 
dead mass, or wood proper, from 
which new tissue is formed; the 
true growing part of plants 
which live several years and at- 
tain a large size. 

of viewing man and nature in- 
augurated in modern times by 
Rene Descartes. The most dis- 
tinctive thing about it is the 
sharpness with which the dual- 
ism, or antithesis, between mind 
and matter stands out in it. Its 
great practical importance for 
the present era lies in its genetic 
relationship to psycho-physical 
parallelism in psychology, and to 
all forms of idealism in philos- 

which the protoplasmic portion 
and nucleus constitute a surface 
layer, the inner mass being 
chiefly yolk, that is food ma- 
terial for the future embryo. 
The eggs of most insects and 
crustaceans are of this type. 

CEPHALTZE. The tendency among 
animals for a head to become 
differentiated from the rest of 
the body. 

produced by the organism, 
either in special glands, the 



glands of internal secretion, or 
in general tissues, and carried 
by the blood and lymph over the 
body generally, to influence the 
growth or functioning of other 
tissues. Hormones is another 
name given to these substances. 

CHEMOTROPIC. Pertaining to the 
reaction of organisms to chem- 
ical stimuli. 

CHROMATIX. The finely granular 
substance most distinctive of the 
cell-nucleus. Its name comes 
from the readiness with which 
it is colored by many dye-stuffs. 

CHROMATOPHORES. Pigment-bear- 
ing sacs, often single cells, in 
plants and animals. It is by 
means of these that the rapid 
color changes in the skin of 
many animals are accomplished. 

CHROMIDOSOMES. One of the many 
names given to minute specially 
stainable bodies in the cytoplasm 
of many cells. 

CHROMOSOMES. The more or less 
definite bodies of the cell-nu- 
cleus into which the chromatin 
granules are grouped. Their 
constancy of structure and rela- 
tion to hereditary characters 
have given them great promi- 
nence in much of recent biolog- 
ical theory. 

CHROMOPHIL. Any body in the liv- 
ing organism that has an avidity 
for staining reagents. 

Coccus. In the classification of 
bacteria according to their 
shapes, those which are spherical 
are called cocci. 

COELEXTERATA. A gl'OUp of aill- 

mals that have a digestive cav- 
ity, but nothing corresponding 
to the abdominal cavity; also 
called radiata from the radial 
arrangement of the body. 

CORTICAL. Pertaining to the cor- 
tex, or outer layer of an organ, 
as of the brain, bark of the tree, 

CRETIX. An individual affected 

with cretinous disease, a disease 
characterized by certain bodily 
deformities and mental impair- 
ments. The malady frequently 
accompanies goitre, and is now 
considered due to deficiency in 
secretion of the thyroid gland. 

CRUSTACEA. A class of inverte- 
brate animals belonging to the 
great phylum arthropoda, briefly 
characterized by their exoskele- 
ton and paired jointed appen- 

CYTOLOGY. Science of the cell. 

CYTOPLASM. Substance of the cell- 
body, as opposed to the cell-nu- 

DETERMIXAXTS. That particular 
class of imaginary ultimate vital 
units by which the development 
of hereditary attributes is deter- 

mined. They were invented by 
Weismann, and were conceived 
to constitute the germ-plasm, 
and to be located primarily in 
the chromosomes of the egg and 
sperm. In later speculation de- 
terminer is used more frequently 
than determinant on some ac- 
count that is not clear. 

DIATOMS. An immense group of 
aquatic, unicellular algae espe- 
cially characterized by their 
firm, box-like, regularly-shaped, 
chitinous shell. 

ganisms which are two-lashed, 
owing to the two flagella pos- 
sessed by most of the species. 
A group of aquatic unicellular 
organisms almost as numerous as 
the diatoms. The photo-synthe- 
sizing power of living substance 
possessed by these two groups, 
and their enormous abundance 
at and near the surface of the 
bodies of water in which they 
live, make them fundamentally 
important for all the life of the 
waters of the earth. 

DISTAL. A common anatomical 
term signifying away from a 



given point, usually some defi- 
nite feature, as the attachment 
of a muscle, taken as a point of 

DORSO- VENTRAL. A term much 
used in the anatomy of th2 
higher animals to signify a di- 
rection from-back-to-belly of a 

DUODENUM. The first portion of 
the small intestine between tha 
stomach and the jejunum. 

DYXAMIC CENTER. A phrase used 
rather frequently in recent biol- 
ogy, especially in the biology of 
the cell, to express the concep- 
tion that certain structures, as 
the centrosome, are in someway 
not clearly specifiable the "seat" 
of various vital activities. The 
phrase has some such implica- 
tion for general physiology as 
"nerve center" had and with 
many still has, for nerve phys- 

ECOLOGICAL. Pertaining to Ecol- 
ogy, the science of organisms in 
relation to their natural environ- 
ments. This old but newly ap- 
preciated and named branch of 
the science of living nature, may 
properly be regarded as the nat- 
ural history of plants and ani- 
mals modified to meet the mod- 
ern demands of comprehensive- 
ness and exactness in dealing 
with a great province of nat- 
ural phenomena. 

ECTOPLASM. The outermost, some- 
what denser layer of protoplasm 
in many cells, especially in many 
unicellular animals. Opposed to 
endoplasm, the inner, more fluid 
mass. The ectoplasm in proto- 
zoans corresponds to the skin of 
higher animals. The presence of 
a more or less sharply set-off 
outer layer or membrane or skin 
in all organisms whatever is 
coming to be recognized as hav- 
ing a more fundamental physio- 
logical meaning than that of a 

protection for the delicate parts 
underneath, now that so much is 

. being learned by physical chem- 
istry about surface phenomena. 

ENERGY. Work and capacity to do 
work. It is important to note 
that work and capacity to work 
necessarily imply some object, 
organic or inorganic, to do the 
work, and hence that when the 
energy of a horse, or of a stream 
of water, is spoken of the word 
energy has a very different 
meaning from what it has in 
such a phrase as the "energy 
conception" of nature or of the 
organism, the implication in 
these cases usually being that 
energy is the real essence of 
nature and of the organism, the 
shape and other so-called static 
attributes which all bodies pre- 
sent, being only incidental and 
mere appearances. 

EXZYME. A chemical substance 
produced by an organism, plant 
or animal, to the end of bring- 
ing about chemical transforma- 
tion in other substances, but 
without itself being transformed. 
The ptyalin of saliva by which 
starch is changed into sugar, is 
typical. Enzymes play a very 
great part, especially in diges- 
tion and nutrition, in the physio- 
logical processes of all organ- 

EPIGEXESIS. That theory of devel- 
opment of the individual organ- 
ism which holds the organs and 
parts to be actually new produc- 
tions, and not merely enlarge- 
ments or actualizations of what 
already existed, this latter con- 
ception of development consti- 
tuting the theory of preforma- 
tion. Although these opposing 
theories were debated with fury, 
almost, some years ago, little is 
heard about them now though 
none of the particular problems 
around which the discussions 



centered can be said to have 
been solved. 

EPIMORPHOSIS. The mode of re- 
generation of organisms in 
which a multiplication of cells 
on the surface of injury is first 
produced, then from this "em- 
bryonal tissue," the new organ 
or part is formed; contrasting 
with morphalaxis, a mode of 
new formation which consists in 
a direct transformation of an 
already existing part into the 
new part. 

FACTOR (in Genetics). A hypo- 
thetical unit of structure or of 
chemical composition, contained 
in the germ-cell, which in some 
way is held to condition the de- 
velopment of a particular char- 
acter in the adult, or of a com- 
plex of characters which are 
transmitted in constant associa- 
tion with one another. Factors 
are believed to interact with one 
another in development, and at 
times to be so "linked" that they 
are only partially independent in 

FLAGELLUM. A lash-like appen- 
dage or large cilium serving as 
an organ of locomotion in some 
Protozoa and some bacteria. 

FORAMINIFERA. A class of rhizo- 
podous marine Protozoans, usu- 
ally having a porous shell. 

substances which are supposed 
to be formed in one part of an 
organism and transported to an- 
other part, there to produce, or 
to influence the production of 
new organs. For example, sev- 
eral botanists have supposed that 
the flower substance of some 
plants is actually produced in 
the leaves. 

GAMETE. A reproductive cell 
which unites with another repro- 
ductive cell to form a zygote. 

GASTRULA. That stage of embry- 
onic development in many ani- 

mals which consists of two germ- 
layers inclosing a central cavity. 
It* is produced from the blastula 
(which see) by the in-sinking of 
one-half of this into the other. 

GEMMIPAROUS. Producing gemma?, 
or buds (reproducing by bud- 
ding), applicable to both plants 
and many animals. 

GEMMULE. 'in the original and 
proper sense a small aggregation 
of cells set apart in the tissues 
of some plants and animals, 
notably in many sponges, for the 
purpose of reproduction. In ori- 
gin and structure gemmules are 
more like buds than eggs, 
though the end served is very 
similar to that served by seeds. 
In a secondary and wholly hypo- 
thetical sense, gemmules are 
imaginary, minute bodies given 
off by all the tissue cells of an 
organism and assembled in the 
germ cells, there to cause the 
development of the next genera- 
tion. This taking of a very con- 
crete name from botany and 
zoology, and using it in a wholly 
imaginary way to explain hered- 
itary development was due orig- 
inally to Charles Darwin, but 
with more or less unimportant 
variations of meaning has since 
been resorted to by many of the 
best known biologists. 
This example indicates the great 
importance for biology, espe- 
cially for the biology of repro- 
duction and development, of dis- 
tinguishing between the same 
terms used in a strictly objective 
and descriptive sense on the one 
hand, and in hypothetical, or 
purely imaginary sense on the 

GENE. A term much used in pres- 
ent-day genetical science, but ap- 
parently not differing in any sig- 
nificant particular from factor 
(which see). 

GENETIC. Pertaining to genetics, 



evolutionary science dealing with 
natural propagation and devel- 
opment, the interest centering at 
present in that portion of devel- 
opment which is hereditary and 
involves sex cells. 

GERM-PLASM. Actually all the pro- 
toplasm of the germ-cells which 
participates in development; the- 
oretically merely the small por- 
tion of the germ-cells supposed 
to be "hereditary substance." 

GONAD. A mass of undifferen- 
tiated, generative tissue from 
which the male and female re- 
productive glands originate. 

GONOPHORE. The ultimate gener- 
ative zooid of a hydrozoan, giv- 
ing origin directly to the genera- 
tive elements. 

HECTOCOTYLIZED. Applied to the 
remarkably altered condition as- 
sumed by one of the arms of the 
male cephalopod to make it an 
organ for impregnating the feT 

HELIOTROPIC. Responding to the 
stimulus of sunlight. 

generation in which the part pro- 
duced is different from that 
which was lost, as, for example, 
when an antenna-like structure 
grows in the place of an eye- 
stalk, in some crustaceans when 
the eye stalk is cut off. 

HISTOGEXESIS. The process of tis- 
sue genesis, or production, from 
undifferentiated cell masses, in 
plants and animals. 

HISTOLOGY. The science of tissues, 
plant or animal; microscopical 

HORMOXE. Literally something 
which excites or stirs up. Orig- 
inally and strictly applied to 
those internal secretions (which 
see), the office of which is to 
incite the parts on which they 
act to greater activity. But in- 
ternal secretions are 'also known 
now which retard or inhibit the 

action of the part they affect; 
and to these it has been proposed 
to apply the term chalone, that 
which slackens. But some phys- 
iologists use hormones as syn- 
onymous with "internal secre- 

HOMONYMOUS. As used in this 
book, an anatomical term refer- 
ring to the different members of 
a series which differ more or 
less, but still all have the same 
general name. Thus all the 
pairs of appendages of a lobster 
are homonymous, or ambulatory 
appendages originally, although 
used for a variety of purposes 

HYDRAXTH. One of the bud-pro- 
duced polyps of a hydroid col- 

IDIOPLASM. Literally plasm which 
is very specially one's own. First 
used to designate the hypothet- 
ical part of the germ-cells which 
is supposed to be alone respon- 
sible for hereditary transmission. 
Idioplasm may be regarded as 
the historical antecedent of 
germ-plasm (which see). 

INTERSTITIAL. Pertaining to or sit- 
uated in an intervening space; a 
term much used in anatomy to 
signify within an organ. 

has long been used in the phys- 
iology of the higher animals, in 
contradistinction to "external se- 
cretion," to designate the prod- 
ucts of glands, like the thyroid, 
which discharge their products 
into the blood or lymph, instead 
of upon the surface of the body 
or into the digestive or some 
other cavity of the body. The 
existence of internal secretions 
was known long before anything 
was known about their use; 
hence this non-committal name, 
so far as function is concerned. 
The recent discovery of their 
office has suggested the name 



hormone (which see) for them, 
and has revealed their great im- 
portance not only for physiology, 
but for philosophical biology. 

INVOLUTION. Literally inrolling, or 
inwrapping. In descriptive biol- 
ogy used to signify the return of 
an organ to its original or nor- 
mal condition after some violent 
or pronounced deformation of it. 
Sometimes, but apparently un- 
justifiably, used as a synonym of 
degeneration. Since the doctrine 
of evolution has become prom- 
inent in biology, a process the 
opposite of evolution has been 
thought by some to be necessary, 
and to this involution has been 

JELLY-FISH. In the interest of dis- 
criminative knowledge, the habit, 
rather common among people 
who have the opportunity to see 
the transparent, somewhat gela- 
tinous-appearing animals of the 
ocean, of calling them all "jelly- 
fishes" should be abandoned. The 
name should be restricted to the 
regularly disc- or dome-shaped, 
tentaculated animals belonging 
to the coelenterate phylum, thus 
enlarging the bounds of definite, 
popular zoological information, 
by recognizing that marine ani- 
mals of several large and very 
distinct classes have this general 
consistency and appearance. 

KARYOPLASM. A cytological name 
referring to the substance or 
plasm distinctive of the cell-nu- 

KARYOSOME. A small, discrete, 
rather constant body which 
stains readily, contained in the 
cell-nucleus; frequently synony- 
mous with nucleolus. 

KATABOLISM. The down-breaking, 
or descensive phase of metabol- 
ism; the opposite of anabolism 
(which see). 

KINETO-NTJCLEUS. One of the nu- 
clei in the two-nuclear protozoa 

supposed to be concerned in 
some special way with the move- 
ment of the flagella or cilia of 
these animals. 

LAMELLAE, singular lamella. A 
term much used in anatomy to 
designate the thin plates, scales, 
etc., that are so numerous and 
varied in form and size in nearly 
all organisms. 

LARVA. Properly applied only to 
stages in the lives of individual 
animals which pass into succeed- 
ing stages through a deep-sealed 
metamorphosis, as for example 
the grub or maggot of a fly, and 
its transformation into the adult. 
Larval stages and profound 
metamorphoses are very common 
and widespread in the animal 

LIMULUS. The technical genus 
name for the horse-shoe crab, an 
animal of special interest to gen- 
eral zoology in several ways. 

MACRONUCLEUS. In the infusoria, a 
group of protozoans, there is one 
large nucleus and one or several 
much smaller nuclei. The first is 
called, from its relatively large 
size, the macro-nucleus; the 
others micro-nuclei. From the 
behavior of the two kinds of nu- 
clei at conjugation and division, 
the micronuclei are known to be 
intimately connected with these 
processes, while the macronucleus 
seems to be more concerned with 
the nutritive functions of the 

MAXUBRIUM. In morphology a 
part or organ which resembles a 
handle; specially the clapper- 
like, or handle-like portion of a 
medusa which is found within 
the "bell." The animal's mouth 
is at the end of the manubrium, 
and most of its digestive cavity 
within the stalk of the manu- 

MATRIX. In biology the ground 
substance in which cells are em- 



bedded in some tissues, and 
which is produced as a secretion 
by the cells. It is one kind of 
intercellular substance. The 
opalescent, almost homogeneous 
chief mass of ordinary cartilage 
is a typical matrix. 

MELANIN. A rather general term 
in biology, especially in zoology, 
applied to dark brown to black 

MEROTOMY. The automatic cutting 
off of parts or segments in liv- 
ing organisms. 

MESENCHYME. Undifferentiated 
mesoderm that produces con- 
nective tissues, some muscles, 
and certain other structures in 
the animal body. 

METABOLIC. Pertaining to metab- 
olism, the process of chemical 
building up and breaking down 
in the living organism. 

METAMERIC. Pertaining to the 
longitudinal series of parts or 
joints into which the bodies of 
many higher animals, such as 
earthworms, lobsters and fishes, 
are divided. 

METAPLASTIC. Pertaining to meta- 
plcMtm applied to changes which 
cells sometimes undergo from 
one plasmic type to another; also 
applied to certain supposedly 
lifeless inclusions in the proto- 
plasm of cells. 

METAZOA. Multicellular animals. 

MICRONUCLEUS. See macronucleus. 

MICROPHYLE. In botany and zool- 
ogy the aperture in the coats of 
the ovule and ovum through 
which the male fertilizing cell 

MONERA. Hypothetical simple 
structureless masses of proto- 
plasm (without any nucleus). 
Assumed by Haeckel as the low- 
est members of the evolutionary 
series. Advance of knowledge 
has found no evidence of such 

MORPHALLAXIS. A kind of regen- 

eration in which part of an or- 
ganism transforms directly into 
a new and different part. 

MORPHOLOGICAL. Pertaining to 
morphology, the science of form 
and structure. 

MORULA. A stage in the embryonic 
development of many animals, in 
which the ovum has completely 
segmented, but the segmentation 
cavity has not yet been formed. 

MYONEME. A thread-like contrac- 
tile structure in the cytoplasm of 
certain higher protozoa. 

NEMATOPHORE. A body of defense 
and offense developed in certain 
hydroids, consisting of a chitin- 
ous receptacle in which thread- 
cells are immersed; the nettling 
organs on the tentacles of large 
j elly-fishes. 

NEURAL. Pertaining to nerves. 

NEUROBLASTS. Undeveloped nerve 

NUCLEO-PLASM. Nuclear substance, 
including the different nuclear 

NUCLEO-PROTEIN. One of the com- 
pounds of nucleins and paranu- 

(EDEMA. Dropsy, a vasomotor 
neurosis characterized by non- 
inflammatory swellings on vari- 
ous parts of the body. 

ONTOGENY. The development of an 
individual organism from germ 
to completed or adult stage. 

ORGANELLE. A little organ, and 
organoid, organ-like, are terms 
applied to the organs of unicel- 
lular plants and animals, not so 
much because of their small size 
and indefiniteness of form and 
structure as on account of the 
theory that a true organ must 
be composed of cells, and cannot 
be a part of a cell. These terms 
are among the sequelae of the 

ORIENTING. Finding or fixing the 
positions or directions. 




terms, basal in Darwin's famous 
hypothesis of heredity, mean all- 
generator and all-generative only 
in the sense that all parts of the 
body of the organism give oif 
gemmules (which see), which 
assemble in the germ-cells to en- 
able these to ?*eproduce the or- 
ganism. Thus the pan, or all- 
generative power was conceived 
as having its original "seat" in 
the organism all-in-all. In other 
words Darwin's speculation was 
almost diametrically opposed to 
the transformation it has under- 
gone latterly, especially in the 
prolific mind of Weismann, the 
germ cells alone, or rather the 
germ-plasm being the all-genera- 
tor, according to these specula- 

PARATHYROIDS. Small glands lying 
near the thyroid but not func- 
tionally connected with the lat- 

.by means of unfertilized eggs. 

PELLICULA. The cuticle or outer- 
most body membrane in some 
unicellular and other low organ- 

PHASE. This old and familiar word 
has taken on new and greater 
importance, both scientific and 
philosophic, with the recent ad- 
vance of knowledge in the region 
of over-lap between physical and 
chemical phenomena, this ad- 
vance making what is generally 
called physical chemistry. A 
phase in pure physics, as it may 
be called, has reference to the 
position of the particles of a 
body when the particles are un- 
dergoing change. For example, 
corresponding particles in two 
succeeding waves of water or air 
are in the same phase. In phys- 
ical chemistry phase has refer- 
ence not to position but to state 
or condition of the constituent 
particles of a heterogeneous, or 

unlike system. Thus, a combina- 
tion of liquid water (in common 
language just water) and solid 
water, or ice, is a two-phase sys- 
tem of water. Philosophically 
viewed, the great significance of 
phases is that the positions and 
states of the particles are possible, 
even conceivable, only in relation 
to the larger, containing part or 
whole. Something of the bear- 
ing of this on the theory of plu- 
ralism (which see), when this 
theory is approached from the 
strictly objective side, will be 
easily seen. 

PHLOGISTON. An imaginary sub- 
stance formerly supposed to ex- 
ist in all combustible bodies, and 
to be the cause of fire and flame. 
For nearly a century before the 
discovery of oxydation as the 
true cause of fire, by Lavoisier, 
the phlogistic theory dominated 
much of chemical science. The 
chief interest in the theory now 
is in its relation to the observa- 
tional and logical processes in- 
volved in interpreting the gen- 
erative processes of nature 
everywhere. The phlogistic the- 
ory may be taken as a type of 
elementalistic causal explanation 
of natural production. 

PHYLOGENIC. Pertaining to phy- 
logeny, the development of the 
race; concerning ancestral or- 
ganisms, real and hypothetical. 

PITUITARY GLAND. A gland of in- 
ternal secretion, situated at the 
base of the brain, and connected 
in the embryo with the roof of 
the mouth. 

PLURALISM, philosophical (so used 
in this book). The conception 
that in its deepest nature the 
universe is multiform and com- 
plex; the opposite of Monism, 
the conception that some single 
Essence or Substance, more or 
less known or unknown, is the 
foundation of all things. 



PLUTEUS. Name given the charac- 
teristic process-bearing larva of 
sea-urchins and their near rela- 
tives. These larvae are of con- 
siderable general interest because 
of the extensive use made of sea- 
urchin eggs in experimental em- 
bryology, the eggs being easily 
obtained and easily kept in the 

PROTEINS. Nitrogenous substances 
found in the bodies of plants 
and animals. These substances 
are usually considered to be the 
most fundamental, from the 
chemical standpoint, in organic 

PROTISTA. A group name intended 
to include all unicellular organ- 
isms; i.e., both protophyta, one- 
celled plants, and protozoa, or 
one-celled animals. 

PSEUDOPODIA. Literally false feet. 
They are temporary protrusions 
of the protoplasm of some pro- 
tozoa, especially of the rhizopo- 
dous class, typified by the amoe- 
ba, the name having reference to 
the locomotor office of the proc- 
esses. But their food-taking and 
digesting office should be noted 

PTYALIN. The unorganized fer- 
ment, or enzyme of saliva, chiefly 
instrumental in the conversion of 
starch into sugar. 

RADIOLARIA. One of the main sub- 
divisions of the protozoa, espe- 
cially characterized by their gen- 
erally spherical outline, and ra- 
diating structures, some soft and 
extensile, others stiff and per- 
manent. The radiolaria are al- 
most all marine. 

REGULATION. Much used in studies 
in the regeneration of organisms, 
to express the power many 
plants and animals have of un- 
dergoing structural or func- 
tional readjustments in order to 
retain, or to regain, their typical 
form; a significant adaptation of 

a general term to a technical 

RETICULAR. Net-like, a term much 
used in anatomy, as many por- 
tions in both plants and animals 
of many grades, present this 
type of structure, though the 
netting never has the regularity 
of manufactured netting. 

RHIZOPODA. The great subdivision 
of the protozoa especially char- 
acterized by sending out pseudo- 
podia (which see). Amoeba is 
usually mentioned as the type of 
this subdivision, but the larger 
number, probably, of rhizopods 
possess shells of one sort and an- 
other, while amoeba is entirely 
naked during all its active life. 

SARCODE. Literally like flesh. The 
name originally applied to what, 
under microscopic examination, 
seemed to be the fundamental 
living-substance of animals. La- 
ter discovered to correspond to 
what was known as protoplasm 
in the cells of plants. 

SARCODYCTIUM. A protoplasmic 
network of the surface of the 
calymma of a radiolarian. 

LATION, &c. It is not without 
philosophical significance that 
the term self has forced its way 
into technical biology, something 
as it has into technical philoso- 
phy. In biology the term is par- 
ticularly common in connection 
with developmental phenomena 
and has reference to operations 
which depend primarily on the 
organism itself, and can be re- 
ferred to "external factors" only 
remotely and in a round-about 

SERICTERIES. Glands by which silk 
and silk-like substances are se- 
creted in many insects. 

SOMA, SOMATIC. The body and 
pertaining to the body. Much 
used in later discussions of 
heredity in a strongly hypothet- 



ical sense, to indicate the com- 
plete independence, so far as 
development is concerned, of the 
body from the germ. The an- 
tithesis is often made stronger by 
speaking of the substance of the 
body and the substance of the 
germ, using the terms somatic- 
plasm and germplasm. From 
the philosophical standpoint it is 
instructive to compare the the- 
oretically complete separation of 
body and germ in modern gen- 
etics, with the theoretically com- 
plete separation of body and 
soul in philosophy and psychol- 

SPECIFICITY. The state of being 
specific, that is, of being mani- 
fested as phenomena distinguish- 
able from all other phenomena. 
The group of terms kindred to 
specific and species, long import- 
ant in systematic and taxonomic 
biology, are becoming increas- 
ingly so with the advance of 
knowledge, especially in the do- 
mains of the chemistry of differ- 
ent kinds of organisms, and of 
comparative behavior and psy- 

SPORAZOA. One of the main sub- 
divisions of the protozoa a lead- 
ing characteristic of which is in- 
dicated by the name, that char- 
acteristic being the commonness 
with which propagation occurs in 
the group by means of spores 
produced within the body of the 
animal. By far the greater num- 
ber of the species of the group 
are parasitic, many of them dis- 
ease producing. 

SPORULATION. The process of con- 
verting into spores, as in the 
sporazoa, in some other animals, 
and in many plants. Spores dif- 
fer from eggs, on the one hand, 
and seeds on the other, only in 
the fact that spores are not sex 
cells, that is, do not need to unite 
with other cells in order to de- 

velop, as is the case with most 
eggs and seeds. 

difference between "spontaneous" 
in this phrase and in the phrase 
"spontaneous action," as of an 
animal, should not be missed. In 
the latter connection the word 
has nearly if not quite its orig- 
inal meaning, that is, of one's 
own accord, or initiative; acting 
by and through one's self alone. 
The adjective pronouns mea, tun 
and siid are said to have been 
used always with sponte in good 
Latin prose. Strictly, then, if 
life really originated from some- 
thing which was not living, that 
is by a "fortuitous" concourse or 
interaction among chemical ele- 
ments of different sort, spon- 
taneous would not be the proper 
term to describe the operation, 
simply because it would have in- 
volved fundamentally several 
selves, even if the different ele- 
ments could each be called a 
self. It would not have been an 
operation identifiable by my, 
your, or his or its, but by their. 
Plurality rather than singularity 
of action would be the essence 
of the conception. 

STEREOTROPIC. Reacting to stimuli 
of contact with solid objects. 

STOLON. A prolongation of the 
body of some plants and animals 
that gives rise to new individuals 
by budding. 

SUDORIPAROUS. Sweat-producing. 

SYMBIOTIC. Pertaining to a state 
of living together of two dissim- 
ilar organisms to the advantage 
of both. 

SYNAPTTC. In cytology pertaining 
to sy nap sis, the conjugation of 
chromosomes in sex cells preced- 
ing the reduction divisions con- 
nected with the maturation of 
germ cells. 

SYNCYTIUM. As used in this book, 
a cytological term applied to a 



protoplasmic mass containing 
many nuclei, but not set off into 
distinct cells. The entire em- 
bryo is of this character in some 
animals. In fact a few embry- 
ologists have contended that dur- 
ing the embryonal stages of 
most, if not all animals, the cells 
are connected by protoplasmic 
strands and bridges, making 
them syncytia. The undoubted 
wide prevalence of syncytial 
structure among animals espe- 
cially, has been used as an argu- 
ment against the cell-theory. 
SYNTHESIS. From the organismal 
standpoint not many terms used 
in biology are more important 
than this. The etymological 
meaning, placed or put together, 
expresses only a part of the to- 
getherness of an organism; the 
part, namely, which pertains to 
the assimilative activity per- 
formed by the organism on its 
nutritive substances. This proc- 
ess may be regarded as a syn- 
thesizing one in nearly the lit- 
eral sense (though even here the 
process is more one of self-ac- 
tivity and less one of external 
agency than seems to be implied 
in the original word). But it is 
when we come to consider the 
original nature and power of the 
organism by virtue of which it 
assimilates food, that the inad- 
equacy of synthesis, except in a 
much modified sense, comes to 
light, for the organism's ability 
to assimilate, that is to put or 
place together, its nutritive sub- 
stances is wholly dependent, so 
far as we have evidence, on the 
fact of its being already and 
originally a together entity. An 
organism is able to put together, 
or synthesize, its food just be- 
cause it itself is together, or syn- 
thesized. A synthesized state is 
a prior condition to synthesizing. 
To be an organism at all is to be 


SYSTEMATIC. Pertaining to a sys- 
tem; literally a standing or being 
together. It is unfortunate that 
"systematic" has come to be re- 
stricted in its application in re- 
cent biology to the formal classi- 
fication of plant and animal 
species. As a matter of fact a 
necessary consequence of the 
unity of all phenomena of the 
living world is that all these 
phenomena "stand" in some nat- 
ural and ascertainable relation 
with all other phenomena, so 
that all biological knowledge 
whatever must of necessity be 
systematic if it really corre- 
sponds to nature. 

TAXONOMY. Mode of arrangement, 
the branch of biology which 
deals with the classification of 
the species of plants and ani- 

TEST. As used in zoology and bot- 
any, an external covering or 
tunic, usually nearly lifeless, 
tough and resistant. Its office is 
mostly protective. 

THYMUS. A gland of internal se- 
cretion found in the neck region 
in all vertebrates, and connected 
originally with the gill system. 

THYROID. One of the most impor- 
tant glands of internal secretion, 
located, as is the thymus, in the 
neck region, but connected em- 
bryonically with the pharynx 
rather than with the gills proper. 

TROCHOPHORE. A larval stage in 
the lives of many marine worms 
and molluscs, characterized by 
being well organized for swim- 
ming by means of cilia variously 
disposed on the surface of the 

TRYPSIN. One of the chief "active 
principles," or enzymes of pan- 
creatic juice. It splits proteids 
into simpler compounds. It is 
produced by some plants as well 
as many other animals than man 



and vertebrates. 

Tuxic. In botany and zoology, 
any well differentiated membran- 
ous covering of an organ or an 
entire organism; much the same 
as a test. 

TUNICATE. Name of a group of 
marine animals, most sharply 
characterized by the cellulose- 
containing tunic, or test, which 
envelopes the body; by the pe- 
culiar basket-like respiratory sys- 
tem; and by the notochord or 
precursor of the vertebral col- 
umn, possessed by all the species 
in the embryonal life, and bj^ a 
few during the whole life, fre- 
quently used synonymously with 
Ascidian, which see. 

VAGAL. Pertaining to the vagus 
nerves, one of the tenth pair of 
cranial nerves in all true verte- 

VASO-COXSTRICTOR. Applied to the 
nerves which cause contraction 
of the walls of blood vessels. 

VASO-DILATOR. Applied to nerves 
which % cause, or more exactly, 
permit a widening of the blood 

vessels by diminishing the tonus 
of the muscles of the vessel 
walls. Since the smaller blood 
vessels are all supplied with both 
constrictor and dilator nerves 
the constant balancing between 
these antagonistic influences, 
both kinds of impulse being in 
response to the general needs of 
the organism, this scheme illus- 
trates well a principle of equili- 
bration widely operative in the 
animal kingdom. 

VISCERAL. A term used in zoology 
to indicate not only the totality 
of internal organs, but also the 
side of the animal on which 
these are situated. 

ZYGOTE. A body formed by the 
conjugation of two reproductive 
cells, called gametes. Gametes 
and zygotes may be either uni- 
cellular organisms, or the repro- 
ductive cells of multicellular or- 

ZYMOGEX. The enzyme-producing 
substance in the secretory cells 

of glands the secretions of which 
contain enzymes. 


Abderhalden, E., i, 102 

Absolutism, metaphysical, ii, 151 

Acacia, ii, 99 

Accidental products of chemical 
change, i, 111 

Acorns, storing of, by woodpeck- 
ers, ii, 269 

Acromegaly, ii, 124, 137 

Action, autocatalytic, ii, 107 

Action-system, ii, 216 

Activity, adaptive, excessiveness 
of, ii, 257 

Activities, instinctive, tendency to 
excessiveness of, ii, 256, 268 

Adaptation, of neural activity, ii, 

Adaptive parallelism, i, 335 

Adaptiveness, ii, 241; of subra- 
tional psychic activities, 250 

Adrenal glands, and nervous sys- 
tem, ii, 131 

Adrenaline, chemical composition 
of, ii, 123 

Adrenin, in blood, ii, 131 ; effects 
on fatigued muscles, 181 

Adsorption, meaning of, ii, 344 

Adventitious buds, i, 38 

Aggregation, principle of, i, 182; 
of protozoa, 268; and synthesis, 
ii, 236 

Air, in relation to consciousness, ii, 
290; breath, and spirits, 338 

Albumen, living and dead, distinc- 
tion between, i, 78 

Alchemy, ii, 288 

Alcyonaria, ii, 97 

Aldrich, T. B., ii, 123 

Alga, i, 41 

Allen, B. M., on origin of sex- 
cells, i, 73; on removal of thy- 
roid, ii, 145 

Alternation of generations, i, 316 

American biologists and organis- 
mal theory, i, 11 

Amoeba, complexity of, i, 289; to 
man, 291 ; as organism and as 
cell, 297 

Amours, of fishes, ii, 266 

Amphibia, i, 204 

Amphioxus, on isolated blasto- 
meres of, i, 204; on graded 
series in, ii, 106 

Analysis, in biological reasoning, 
ii, 206; and abstraction, 234; and 
synthesis, 235; remarks on, 236; 
of organic substance, 341 

Anatomist, of protozoa, i, 286 

Ancestral, i, 290 

Anger, ii, 132, 317 

Animal, odors of, i, 84; behavior, 
ii, 208; human, 284; kingdom, 

Animals, as analytical chemists of 
one another, i, 88 

Annelids, ii, 192 

Antagonisms, ii, 175; within au- 
tonomic nervous system, 178; co- 
operative, 178 

Anthropologist, ii, 285 

Anthropology, ii, 227 

Anthropomorphism, ii, 201 

Anti-bodies, i, 100 

Anticipatoriness, ii, 242 

Antigens, i, 100 

Ants, behavior of, ii, 257, 279; 
larval, spinning of cocoon by, 

Aphids, i, 353 

Appendicularia, ii, 3 

Apperception, relation to tropisms, 
ii, 221 ; definition by Wundt, 
233; real nature of, 243 

Apples, odor of, i, 87 

Argon, ii, 341 

Aristotle, i, 2; ii, 278 

Arrhenius, S., i, 101 

Art, creativeness in, ii, 223; crea- 
tive impulse in, 227 




Artificial parthenogenesis, i, 345 

Ascidian, bud propagation in, i, 
50, 309; egg development in, ii, 
17; tentacles of, 98; ganglion 
of, 187 

Assimilation, ii, 205 

Association in psychology, objec- 
tive and subjective sides of, ii, 

Associationism, ii, 228 

Associationists, ii, 221 ; and ele- 
mentalists, 228 

Atomistic theory, ii, 151 

Atoms, ii, 149, 160 

Attention and choice, ii, 231 

Attitudes, difference between sci- 
entific and philosophic, ii, 307; 
elementalist, and emotions, 321 

Attributes, correlation of, i, 215; 
ii, 202; physical or material, and 
psychical or spiritual, 215; struc- 
tural and functional, 277; eth- 
ical, 284; observed corporeal, 
289; physical and chemical, 305; 
latent of oxygen, 341 ; latent, 

Autocatalytic action, factor in 
growth, ii, 107 

Automaticity, ii, 221 

Autonomic nervous system, ii, 128; 
vagal or cranial, 129; sympa- 
thetic or thoracico-lumbar, 130; 
sacral, 130; antagonisms within, 

Avoiding reactions, ii, 252 

Axioms, ii, 297 

Axones, ii, 170 

Baboon, i, 98 

Bacillus butschlii, i, 262 

Bacteria, i, 310; membrane and 
surface structure, 257; un- 
doubted organisms, 263; classifi- 
cation of, 266 

Balance of organs, i, 7 

Balanoglossus, i, 223 

Balfour, F. M., i, 268 

Ballowitz, E., ii, 2 

Barker, L. F., on interrelation be- 
tween internal secretions and 
nervous system, ii, 130, 138 

Basedow's disease, ii, 138 

Bateson, W., i, 22 

Bayliss, on accidental chemical 
products, i, 112; and Starling, 
ii, 120; on meaning of autonomic 
nervous system, 129 

Bearers, i, 306; of heredity, 338 

Bees, honey, ii, 268 

Behavior, complexity of, i, 289; 
animal, ii, 208, 210, 227; be- 
havior-knowledge, 277 

Behring Sea, ii, 211 

Benda, C., ii, 35 

Benecke, W., i, 258 

Bernard, Claude, ii, 149 

Bichat, i, 4 

Bio-chemical substances, phylog- 
eny of, i, 110 

Bio-chemistry, and taxonomist, i, 

Biococcus, i, 22; 319 

Biogen conception, i, 194 

Biogenesis, theory of, i, 27; versus 
spontaneous generation, 316 

Bio-integration, types of, ii, 94 

Biologist, anthropological, ii, 335 

Biology, real, i, 5; goal of, ii, 
152; elementalist, and associa- 
tionist psychology, 228; subdivi- 
sions of, 283 

Birds, high flight of, ii, 258; song 
habits of, 260; mating habits of, 

Blastomeres, i, 203; position in the 
whole, 206 

Blastula, i, 203 

Blepharoplast, i, 255, 329 

Blood, and bloods, i, 91 ; cor- 
puscles, white, 297; adrenin in, 
ii, 131 

Blue-jay, storing habits of, ii, 270 

Body, i* 321; ii, 150, 215; relation 
to mind, 216; constitution of, 
289; vs. corpse or cadaver, 322; 
and soul, 323 

Born, G., i, 207 

Botanical, diagnosis, i, 265 

Botany, elementary instruction in, 
i, 236 

Boyle, Robert, ii, 288 

Brain, not coordinating center, ii, 
191 ; normality dependent on, 
194; as element of organism, 216 



Brandt, Percy, i, 39 

Bread, "secret powers" of, ii, 300 

"Breath of Life," ii, 303, 336, 


Brown, A. P., i, 95 
Briicke, E., conception of the cell, 

i, 129 
Bud, adventitions in plants, i, 38; 

propagation in compound ascid- 

ian, 50, 309; in bryozoa, 53 
Burrows, M. T., on tissue cultures, 

i, 173; on organ formation in 

such cultures, 176 

Butterflies, larvae of, ii, 239 


Calkins, G. N., i, 240 

Cambium, buds from, i, 39 

Canary birds, ii, 261 

Canidae, i, 96 

Cannon, \V. A., on chromosomes 
and Mendelism, i, 356; on 
heredity in plant hairs, ii, 55 

Cannon, W. B., on autonomic nerve 
action and adrenin, ii, 129, 131, 
162, 178, 185, 319, 323 

Carbon, ii, 338 

Carrel, Alexis, on tissue cultures, 
i, 168, 174; isolated tissues and 
"morphological plan" of organ- 
ism, 177 

Carrying, characters of adult, i, 
224; hereditary qualities, second- 
ary and acquired, ii, 67 

Cassia, ii, 99, 106 

Casteel, D. B., ii, 9 

Castle, W. E., definition of hered- 
ity, i, 315 

Caterpillar, ii, 239 

Caudal tube of spermatozoan, ii, 

Causal factors, ii, 103 

Cause, in heredity, i, 313; sufficient, 
ii, 147; contributing, 153; un- 
known, of experience, 303 

Cell, as chemical laboratory, i, 82; 
physical chemical conception of, 
116; as an organism, Briicke, 
129; as key to ultimate biologi- 
cal problems, 163, 181; in logical 
and factual aspects, 228; as 
elementary organism, 228; ag- 
gregations, 295; colonies, 295; 

evolution of, Minchin, E. A., 

Cell-membrane, produced by pro- 
toplasm, ii, 59 

Cell-nucleus, and protoplasm, i, 

Cell-state, i, 295 

Cell-system, i, 216 

Cell-theory, what it is, i, 150 et 
seq.; inadequacy of, 11 and 158; 
attempt to subordinate protista 
to, 280 et seq.; and integration 
of nervous system, ii, 169 

Cell-wall, in higher plants, ii, 51 

Ceils, subordinate to living beings, 
i, 293; used by living beings, 
294; isolated, 294 

Cellular centers, i, 332 

Centers, dynamic in cells, i, 333; 
of apperception, ii, 233 

Centrioles, i, 333 

Centrosome, i, 330, 331 

Cerebral cortex, ii, 216 

Cerebrum, ii, 130 

Ceremonies, self-exhausting, of 
mating, ii, 264 

Chaetognatha, ii, 281 

Chcpfopterus, i, 12 

Chain reflexes, ii, 197 

Challenger Expedition, ii, 7 

Characters, special and general 
in heredity, ii, 40 

Chemical action and interaction, i, 
215; autocatalytic in organic 
growth, ii, 105 

Chemical basis of genus and spe- 
cies, i, 107 

Chemical, criterion of, ii, 289 

Chemical messengers, i, 23; ii, 119, 
121, 128, 170 

Chemico-functional integration, ii, 

Chemico-naturalist inquiries, i, 105, 

Chemist, and naturalist, i, 107 

Chemistry, and organism, i, 75; of 
organisms, 91; comparative, 105; 
and variation, 115; in solving 
problems of heredity, ii, 42; 
definition of, 287; physical, 303, 
336; periodic law of, 329; atom- 
istic, 343 



Chicken pox, i, 264 

Child, C. M., .and physiological cor- 
relation, i, 17; and metabolic 
gradients, ii, 108 

Chipmunk, storing habit of, ii, 271 

Chondriosome, ii, 36; as material 
substratum of different tissues, 

Chromatin, theory of, i, 314; rela- 
tively undiiferentiated, 318; evi- 
dence of, as hereditary sub- 
stance, 326; physicial basis of 
heredity, 328; supposed omnipo- 
tence in heredity, ii, 14; kinds 
of, 67 

Chromatinists, i, 319 

Chromosomal elementalism, i, 320 

Chromosomal hypothesis of hered- 
ity, evidence for, i, 324, 326 et 

Chromosome dogma, ii, 59 

Chromosomes, i, 21, 306, 324; as 
immediate ancestors, 319; in 
fertilization, 342; accessory, 347; 
X and Y, 350; seat of inheri- 
tance material, ii, 22; in rela- 
tion to heredity, 66; initiators in 
heredity, 83 

Chun, C., i, 301 

Cilia, i, 330 

Ciliary tuft of spirillum, i, 259 

Classification, i, 99, 296; of 
physical facts, ii, 217; synoptic, 

Chemical evidence of adrenal- 
nervous connection, ii, 133 

Cohn, F., on classification of bac- 
teria, 266 

Collins, H. H., ii, 258 

Common-paths in nerve physiol- 
ogy, ii, 171 

Common-sense, i, 32 

Comparison, i, 99; of shells of 
rhizopod and nautilus, 237; 
sacrifice of, in experimental 
method, ii, 279. 

Competition, ii, 175 

Condition, molecular appeals to, 
i, 276 

Conjugation, i, 269 

Conklin, E. G., on egg as stage in 
life of organism, i, 193; on 

development of Ascidian egg, ii, 
17; on hereditary characters de- 
termined by cytoplasm, and by 
chromatin, 42, et seq. 

Coordination, neural, not a "cen- 
tral" process, ii, 192 

Corycella, i, 270 

Consciousness, ii, 161; contents of, 
233; organismal theory of, 282; 
and chemical action, 290; theory 
of, and theory of knowledge, 
296; an attribute of the organ- 
ism as a whole, 309; and 
physico-chemical conception of 
organism, 324; and pro-con- 
sciousness, 350 

Contents, of consciousness, ii, 225, 

Courtship of animals, 262 

Cowdry, E. V., i, 437 

Crampton, H. E., ii, 26 

Crane, sand-hill, ii, 258 

Crepidula, ii, 20 

Cretin, ii, 116 

Crickets, chirping of, ii, 261 

Crithiditi, i, 334 

Ctenophore, i, 201 

dishing, H., ii, 113, 124 

Cuvier, i, 5 

Cycads, ii, 58 

Cytoplasm, and Karyoplasm, i, 
'135; Kinds of, ii, '67; funda- 
mental and primitive as heredi- 
tary substance, 68 

Cytoplasmic activity, in spicule 
production, ii, 52 

Cytoplasmists, i, 319 

Cytostome, i, 248 

Dances, of lapwing, ii, 262 

Darwin, Chas., as naturalist, i, 75; 
as example of creativeness in 
science, ii, 225; on comb of hive 
bee, 168 

Davidson, H. C., on plant as sym- 
biotic colony, i, 35; on "planto- 
gens," 36 

De Bary, and cell theory, i, 162 

Definition, i, 296 

Dendrites, ii, 170 

Descartes, Rene, ii, 298 

Descent, i, 315 



Description, i, 99 

Determinants, Weismannian, i, 21, 
225; changed to determiners, 348 

"Determined," different meanings 
of, ii, 49 

Determiner, fascination of for 
some minds, i, 306; meaning in 
true objective sense, ii, 16; and 
dialectics, 76; theory of, con- 
trary to chemical principles, 
79; Wilson's proposal to drop, 

Development, in protozoa, i, 267; 
cause of, ii, 158 

Developmental mechanics, i, 18 

Dewey, John, ii, 298; on "Self" 
and environment, 305 

Diagnosis, medical, i, 265 

Dialectics, and determiner hy- 
pothesis, ii, 76 

Diatoms, i, 310 

Difference, chemical, between or- 
ganisms, i, 83; between germ- 
cells, importance of, 214; re- 
semblances and, 317; in func- 
tion and behavior, ii, 276 

Differential factor, ii, 82 

Differentiation and integration, ii, 

Dinoflagellates, i, 310 

Directing activity, of develop- 
mental process, i, 70 

Display, mutual, ii, 264 

Distribution, vertical, ii, 281 

Division, of labor, i, 205; deter- 
mined by growth, 220; physio- 
logical, ii, 24 

Dobell, C. C., on nuclei in bac- 
teria, i, 262; on Ehrenberg's 
conception of protozoa, 284; on 
protozoa as non-cellular, 290 

Dog, as causal explanation, ii, 
203, 204 

Donaldson, H. H., ii, 169 

Doncaster, L., i, 352 

Dormitive principle, ii, 204 

Dramatist, ii, 317 

Driesch, Hans, on cell theory, i, 
153; totipotence theory of, 202, 
et seq. 

Ductless glands, ii, 114 

Dujardin, Felix, interpretation of 

protozoa, i, 280; and plasmic 

elementalism, 320 
Duodenal mucous membrane, ii, 

Dynamic center, of cell, i, 333 

Earthworm, ii, 191 

Echinus, i, 202 

Ecology, ii, 212, 279 

Economy, physiological, ii, 261 

Ectoderm, i, 46 

Effect, i, 313 

Egg, of chick, studied chemically, 
C79; of frog, 199; floating, 213; 
hereditary attributes of, 214; as 
stage in development of indi- 
vidual, ii, 24 

Ehrenberg, C. G., interpretation 
of protozoa, i, 280, and ii, 66 

Elements, photosensitive, ii, 189; 
physical and chemical, 235; 
psychical, 235; chemical, cri- 
terion of, 286 

Elementalism, i, 2; narrowing in- 
fluence of, 230; cellular, 286; 
and internal secretion, ii, 141 

Elementalist, conception, i, 280; 
speculation, 319; and organismal 
standpoints, ii, 148; theory, and 
neglect of fact, 157; anarchistic, 
160; attempt to interpret trop- 
istic and segmental theories, 
198; biology and associationist 
psychology, 228 

Elementalistic interpretation, ii, 

Elementary organism, i, 227 

Embryogeny, i, 277 

Embryology, methods of, i, 224; 
of protozoa, 268; and genetics, 
311, 324 

Embryo, i, 204, 272 

Emotions, and physical organiza- 
tion, ii, 216; natural history de- 
scription of, 318, 322; elemen- 
talist description of, 32J 

Emotional, attitude, i, 322; Glyco- 
suria, ii, 132; psychic life of 
animals, 133 

Empedocles, i, 3, 40 

Endoderm, i, 46 

Endocrine glands, ii, 114, 130 



ii, 157; matters of, 299 

Endoplasm, i, 277 

Energy, formative and regenera- Falta, W., ii, 134 
tive, i, 11; of modern physics, Fatalism, ii, 89 
76; and matter, 141; conception Fatigue, and sugar in blood, ii, 

132; and blood pressure, 181 
Fear, and adrenin in blood, ii, 
132; with physical and psychical 

of, 320; of contraction, ii, 63; 

and substance, 337; and power, 

force, work, 342; surface, 344 
Entelechy, ii, 149 
Environmental influence, ii, 244 
Enzymes, digestive, comparative Felidae, i, 96 

chemistry of, i, 104; facilitate Ferns, ii, 99 

transformation, ii, 81 
Epicurus, i, 3 

Epithelium, and internal secre- Finalism, ii, 152 

life, 183 
Feeling, intellect, will, ii, 217 

Fertilization, hybrid, i, 344 
Fielcle, A., on odors in ants, i, 89 

tions, ii, 135 

Eppinger, H., ii, 133 

Equilibrium, general notion of, i, 
17; physical chemistry concep- 
tion of, 216 

Essences, ii, 288 

Esterlv, C. O., ii, 208 

Finch, house, ii, 260 

Fishes, sperm of different species, 
i, 102; floating eggs and young 
of, 213; mating habits of, ii, 265 

Flagella, of bacteria, 259; rela- 
tion to nucleus, 328, 330; struc- 
ture and origin of, 370 

Evidence, direct and indirect, of Flat-worms, ii, 109 

mechanism of heredity, i, 325; Flavors, of animals and plants, i, 
favorable to chromatin as hered- 84 

itary substance, 326; visible and Fluid, as phase of system, i, 216 
invisible, ii, 56; picking out of, Foetus, i, 272 

Forbush, E. H., ii, 270 

ii, 263 

Evolution, i, 291, ii, 241 

Evolutions, four simultaneous, i, 

Eudendrium, i, 67 

Eugenics, i, 305; and fatalism, ii, 

Excessiveness, of instinctive activ- 
ity, ii, 259; of sex impulse, 265 

Forces, antagonistic, ii, 127; con- 
stitutively antagonistic, 134; 
abuse of the term, 298 

Fore-foot, horse's, and man's hand, 
ii, 224 

"Formative stuffs," i, 16; and in- 
ternal secretions, ii, 142; of 
Sachs, 147 

Excitability, threshold of, ii, 165; Form^-determination, cytoplasmic, 

selective, 165 
Excitor, ii, 147 

ii, 14 
Forsyth, Ruth, ii, 44 

Experience, ii, 305; subjective and Fossil wood, ii, 58 

objective, 285; causes of, un- Foster, Sir Michael, ii, 346 
known, 303 Fragmentation, and analysis,, ii, 

Experimentation, limitations of, ii, 236 

212; laboratory, 279 Freud, S., ii, 350 

Explanation, spurious type of, ii, Friedlander, B., ii, 102 

81; causal, 146, 240; of trop- Frog, eggs of, i, 199; pigment 

cells of, 339; sex determination 
in, ii, 76; larva and thyroid of, 
143; croaking of, 261 
Fruit flies, supposed connection 
between mutations and chromo- 

isms, 190 
Expression of emotion, ii, 264 

Factorial hypothesis in heredity, i, 

Factors, in heredity, i, 42, 306 

somes in, i, 354 

Facts, neglect of, by elementalists, Fundamental, criterion of, ii, 201 



Fungus, i, 41 

Fur-seal, development of sperm 

of, ii, 4; migrations of, ii, 210; 

mating habits of, 267 
Fusibility of tissues, as test of 

relationship, i, 143 

Gall-stones, ii, 164 

Gametes, i, 269 

Ganglia, supra-oesophageal, ii, 
191; as relay stations, 194 

Gastrula, i, 203 

Gates, R. R., i, 353 

Gemmule, as method of asexual 
propagation, i, 309; as specula- 
tive entities (see term in glos- 

Gene, a hypothetical entity in 
modern genetics, i, 20; static 
nature of the conception, 42; 
historic antecedent of, ii, 84 

Geneticists, modern school of, i, 

Genetics, tendency to ignore em- 
bryology, i, 311; inattention to 
developmental facts, ii, 14; 
thinking on, 72 

Germ, nature of, i, 223; imaginary 
structureless, 268 

German Deep-Sea Expedition, i, 

Germ-cells, in hydroids, i, 60; 
promorphology of, 211; "throw 
of" the soma, 319; subject to 
metabolism, ii, 74 

Germ-disc, i, 208 

Germinal, continuity, i, 310; ma- 
terial, 209; localization, ii, 16 

Germ-layers, i, 45; subservient to 
organism, 48; and the germ- 
plasm theory of Weismann, 58 

Germ-plasm theory, type effect of, 
on observation, i, 66; implica- 
tions of, 318; extreme form of 
contrary to inductive science, ii, 

Germ-regions, organ forming, i, 

Giardia, i, 254 

Gibbs, J. Willard, ii, 341 

"Gifts," natural, ii, 235 

Ginkgo. i, 330 

Gladness, fits of, ii, 263 

Glands, ductless, endocrine, ii, 114 

Glycogen, i, 215 

Goal of biology, ii, 152 

Goette, Alexander, review and 
criticism by Weismann on sex- 
cells in hydroids, i, 62, et seq.; 
organismal trend of results by, 

Goitre, ii, 115 

Gradients, direct and inverse, ii, 
106; axial metabolic, 107 

Grafts, i, 143 

Granules, epithelial, ii, 135 

Grasshopper, i, 357 

Graves' disease, ii, 133 

Grebe, great crested, ii, 263 

Grennell, Joseph, i, 85 

Groos, Karl, ii, 273 

Growth, determines division, i, 
220; integration, ii, 93, 94; cycle 
of, 105; of an organism, 105; 
explanation of, 107 

Gudernatsch, J. F., ii, 143 

Habits, mating, of birds, ii, 263; 
of viviparous fishes, 265; stor- 
ing, of honey bee, 268, of wood- 
pecker, 269, of mammals, 271; 
in nature, 278 

Haeckel, E., on radiolaria, i, 235; 
moneron theory, 256, 320 

Haecker, V., on radiolaria, i, 236; 
on ontogeny of radiolaria, 278 

Hairs of higher plants, ii, 55 

Half-embryo, i, 199, 201 

Hallez, P.," i, 218 

Hardens, A., i, 104 

Hargitt, C. \V., i, 67 

Harmer, S., i, 54 

Harmonic, equipotential system, 
i, 205; equilibrium, ii, 118 

Harmony, i, 3; in health, ii, 127 

Harrison, R. G., on tissue cul- 
tures, i, 168, et seq. 

Heidenhain, H., i, 332 

Heinke, Fr., i, 213 

Heliotropic, ii, 240 

Helium, ii, 342 

Hemoglobin, i, 95 

Henderson, L. J., ii, 339 

Herbart, as extremist in associa- 



tionist psychology, ii, 229 

Herbst, C.,*i, 344 ' 

"Hereditary substance," contrary 
to facts in hydromedusae, i, 68 

Heredity, and clementalistic phil- 
osophy, i, 20; nature of, 305; 
stronghold of biological elemen- 
talism, 305; effort to restrict to 
sexual propagation, 308; defined 
by E. G. Conklin, 308; com- 
plex of causes, 313; definition, 
314; chromatin in, 320, 321; and 
sex, 348; cytological basis of, 
349; spermatozoon's tail mani- 
festation of, ii, 3; mitochondrial 
theory of, 33; and sponge spic- 
ules, 53; summary of informa- 
tion on physical basis of, 64; 
narrowing definition of, 85 

Herlitzka, Amedeo, i, 205 

Hertwig, O., theory of Biogene- 
sis of, i, 27; experiments on 
half embryos, 200; on centro- 
some, 332 * 

Hertwig, R., on cell-theory as ap- 
plied to protozoa, i, 288; on 
experimental determination of 
sex, ii, 76 

Higher Usefulness of Science, ii, 

His, Wilhelm, i, 208 

Histogenesis, and the mechanism 
of heredity, i, 325, and ii, 32; 
and species characters in adults, 

Hobbes, Thomas, and sensational- 
ism in philosophy, ii, 219 

Holmes, S. J., an organism as 
symbiotic community, i, 183; on 
brainless frogs, ii, 195; on ac- 
tivities of Amphithoe, 248; on 
behavior of ants, 257 

Hooker, D., i, 339 

Hopkins, F. G., on the cell as a 
chemical laboratory, i, 82; on 
the physical chemistry of the 
cell, 114 et seq.; on the cell 
constituents essential to the cell 
as a system of phases, 192; ver- 
sus particular types of mole- 
cules as an explanation of life, 

Honey-bee, eggs and chromosomes 
of, i, 352; storing habits of, ii, 

Hormones, importance of, to or- 
ganismal conception, i, 23; na- 
ture of action of, ii, 121 ; rela- 
tion to nerve action, 128; sup- 
posed identification with "form- 
ative stuffs," 142; integrative 
office of, compared with that of 
nervous system, 162 

Hudson, W. H., ii, 261 

Human being in one-celled stage, 
i, 217 

Hume, David, ii, 298 

Hunter ciliates, i, 235 

Huntsman, A. G., ii, 44 

Huxley, Julian S., ii, 263 

Huxley, T. H., on the physical 
basis of life, i, 121; on the 
cell-theory, 288, 296; on innate 
ideas as conceived by Descartes, 
ii, 298; against materialism, 302 

Hyclroids, germ-cells in, 60; sepa- 
rated blastomeres in eggs of, 
204; graded growth series in, ii, 

Hypophysis, ii, 124; alliance with 
thyroid and adrenals, 127 

Hypopituitarism, ii, 113 

Hypothesis, ii, 282; of conscious- 
ness, 286; "working," 291 

Idea, central of this book, i, 24 
Ideas, atomistic and association 
of, ii, 229; "relations of," in 
Hume's system, 299; innate, 301 
"Identical stuffs," i, 123 
Impertinence, scientific, ii, 247 
Individual, man, i, 31 ; tree, 31 ; ex- 
altation of, 196; normal, ii, 205 
Individuality, in the living world, 
i, 30; Huxlej^an, 43; of chromo- 
somes, 85; of organism, ii, 111 
Inheritance, i, 312; nuclear theory, 
and cytoplasmic localization, ii, 
22; of acquired characters, 24; 
material imitator rather than 
determiner, 66; probability that 
substance becomes such in each 
ontogeny, 73 
Inhibition of reflexes, ii, 176 



Initiative, mental, ii, 244 

Initiator hypothesis in heredity, 
advantages of, ii, 83 

Inner mass, and outer layer, of 
body, i, 301 

Insane-like conduct from absence 
of brain, ii, 194 

Insect, eggs of, i, 220; sperm of, 
ii, 9 

Instinct, involves animal as a 
whole, ii, 188; importance of as 
zoological term; 247; variability 
of, 251; and intelligence, 256; 
food-gathering, 268; problem of, 
284; "instinct actions," "instinct 
feelings," 284; and physical or- 
ganization, 310 

Integratedness, of Giardia, i, 255; 
and equilibrium, ii, 198 

Integration, growth, ii, 93; func- 
tional, 113; distinction between 
developmental and functional, 
161; cellular, in reflex arc, 163; 
and differentiation, 168; psy- 
chic, 214 

Integrity, organismal, i, 26 

Intellect, feeling, will, ii, 217; and 
instinct, 256 

Interaction, chemical, i, 215 

Intercellular substance, ii, 170 

Interdependence, metabolic, ii, 104 

Internal secretions, importance of, 
to organismal conception, i, 23; 
conception of, ii, 113; interrela- 
tions of, 124; and epithelium, 
135; and "formative stuffs," 142 

Internal secretory systems, ii, 128; 
and nervous systems, 128 

Investigation of distribution of 
sex-cells, i, 73; statistical of 
animal behavior, ii, 280 

Iron, ii, 329, 337 

Irradiation, ii, 174 

Isolated cells, i, 167 


James, William, on human energy, 
ii, 132; on associationist psy- 
chology, 229; on consciousness 
of self, 309 

Jelly-fish, i, 235 

Jennings, H. S., on ultra-Mendel- 
ism, i, 42; on complexity of 

protozoan behavior, 289; on trial 

and error, etc., ii, 252 
Johnson, H. P., i, 272 
Jolly, J., i, 176 
Jordan, H. E., ii, 60 

Kant, E., self-conscious unity of 
apperception, and transcenden- 
talism, ii, 233 

Karyoplasm, i, 135 

Kelp, ii, 105 

Key, cell as, to biological phe- 
nomena, i, 229 

King, Miss H. D., ii, 76 

Knowledge, analytic and synthet- 
ic, i, 25; -getting, ii, 213; ana- 
lytic, 213; theory of, 296; nature 
of, 297 

Kofoid, C. A., i, 244; on soil amoe- 
ba, 328 

Kofoid and Christiansen, on neu- 
rometer system in flagellates, i, 

Kolle and Wassonnann, i, 258 

Laboratory, as an agency, ii, 212 
Lamarck, i, 5, 75 
Lancelet, ii, 95 
Langley, J. M., ii, 129 
Lapwing, dance of, ii, 262 
Larvae, grafting together, i, 207; 

specific characters in, 214; as 

"carrier" of adult characters, 


Law, of elements, ii, 160 
Lead, ii, 341 
Leaves, compound, ii, 99 
Leeches, ii, 311 
Leibnitzian theory, ii, 150 
Lemon trees, i, 38 
Lewis and Lewis, ii, 40 
Life, ii, 286; vegetal, emotional, 

rational and intellectual, 162; 

tripod of, 182; subjective, 284 
Like produces like, i, 315 
Likenesses, functional, ii, 276 
Lillie, F. R., as pre-organismalist, 

i, 11; on "properties of whole" 

in the embryo, 12 and 193 
Lillie, R. S., i, 340 
Limulus muscle, ii, 60 
Lineus lacteus, i, 189 



Living beings, in nature, ii, 

Living substance, i, 115 

Living units, hypothetical, i, 19 

Localization, by protoplasmic flow- 
ing, ii, 19 

Locust, i, 218 

Locy, Wm. A., i, 280 

Loeb, Jacques, i, 23; on relative in- 
fluence of nucleus and protoplasm 
on heredity, ii, 41; on identifica- 
tion of internal secretions with 
"formative stuffs," 141; on 
"ultimate aim" of biology, 151; 
and "organism as a whole," 185; 
neglect by, of work of Sher- 
rington and Cannon, 185; on 
understanding of natural phe- 
nomena, 207; organismal ten- 
dency of tropism theory of, 
240 ' 

Logic, pure, i, 22 

Loomis, L. M., i, 85 

Love, emotions of, ii, 265 

Luciani, L., ii, 115 

Lucretius, i, 3 

Machines, living, ii, 252 

Macrocystis pyrifera, ii, 105 

Mammals, storing habits of, ii, 271 

Manly, J. M., on "exuberant vital- 
ity" of Shakespeare, ii, 223 

Marceau, F., ii, 61 

Marshall, F. H. A., ii, 79 

Materialism, author's attitude to- 
ward, ii, 207; Huxley against, 

Mathematics, ii, 297 

Mating habits, of birds, ii, 263; 
of fishes, 265 

Matter, and energy, in modern 
physics, i, 76, 141 ; and force, 
196; composition of, 288, 341; 
generality of the term, 304 

Mass action, ii, 344 

McClung, C. E., i, 347 

Mcllvane, Charles, i, 87 

McMurrich, J., Playfair, on germ 
layers, i, 47; character of cell 
division in embryo, 219 

Meadow lark, western, son-g habit 
of, ii, 259 

Mechanism of heredity, i, 315, 
322; organic vs. inorganic, ii, 252 

Medussetta, i, 236 

Meirowsky, E., i, 339 

Melanin, i, 339; formed in cyto- 
plasm, 341 

Membrane, and surface structure 
of bacteria, i, 257; synaptic, be- 
tween cells of reflex arc, ii, 167 

Mendel, Gregor, i, 305 

Mendelian inheritance and chro- 
mosomes, i, 356 

Mendelism as a creed, i, 324 

Mental, sense, i, 3; initiative and 
restlessness, ii, 243 

Meristic parts, ii, 95; meristic phe- 
nomena in plants and in ani- 
mals, 103 

Merotomy, i, 276 

Mesenchyme cells, dormant in tad- 
poles, ii, 147; as inheritance 
material, 155 

Messengers, chemical, secretin as 
example of, ii, 119, 121 

Mesoderm, i, 46 

Metabolic processes, and the or- 
ganism's supremacy over its 
cells, i, 294; interdependence of, 
ii, 104 

Metabolism, i, 215; germ cells sub- 
ject to, ii, 74; katabolic and 
anabolic, 346 

Metals, "base" and "noble," ii, 288 

Metameres, ii, 95 

Metamorphosis, ii, 145 

Metaphysician, ii, 285 

Metaphysics, chromatin and, i, 
321; juvenile, ii, 141; as an epi- 
thet, 201 

Metaplasy, of differentiated cells, 
i, 186 * 

Metazoa, i, 268 

Metcalf, M. M., i, 289 

Method, experimental, ii, 278; sta- 
tistical, employed at Scripps In- 
stitution, 280; natural history, 
in study of self, 282; impor- 
tance of, 282 

Meves, F., ii, 35 

Meyer, Arthur, i, 258 

Mice, summersaults of, ii, 258 

Michael, E. L., ii, 281 

Microbes, specificity of, i, 265 

iMiescher and Kossel, i, 79, 102 
Migration, of sex-cells, i, 61; ex- 



tent of, by birds, ii, 259 

Milk, i, 104 

Minchin, E. A., Biococcus theory 
of, i, 2; on evolution of cell, 
307; as chromatinist, 318 

Mind, relation to body, ii, 215, 
216; "mind stuff," 3lS 

Miracles, i, 322 

Mitochondria, in ontogeny of in- 
sect sperm, ii, 9; not trans- 
formed into neuro-fibrils, 37 

Mitochondrial theory of heredity, 
ii, 33 

Moeser, W., ii, 107 

Molecular condition, appeals to, i, 
276, 282 

Molecules, ii, 149 

Mollusca, i, 221 

Moore, V. A., i, 259 

Monogamy, in viviparous fishes, 

11, 265 " 

Moneron theory of Haeckel, i, 256, 

Morgan, C. Lloyd, ii, 328 

Morgan, T. H., and regeneration, 
i, 180; on half-embryos of frog, 
200; on eggs of phyiloxerans, 
353; on mutations and heredity 
in fruit flies, 354 

Morphallaxis, i, 180 

Morphological, plan of the organ- 
ism, i, 177, 195; entity, funda- 
mental to tropism conception, ii, 

Mortensen, T. H., i, 216 

Mosaic, theory of development, i, 

12, 198, 205; "mosaic picture," 
organism as, 208 

Mosquitoes, i, 213 
Mosses, leaves of, ii, 57 
Mushrooms, poisonous species of, 

i, 87 

Musk-deer, i, 86 
Mustard plant, weight of seeds of, 

ii, 102 
Mutation, connections of, with 

chromosomes, i, 353 
Myomeres, ii, 95, 106 
Mystification of protoplasm, i, 121 

self, ii, 282; importance of, 283 

Natural selection, and selected 
evidence, ii, 263; slight regard 
for quantity by, 259 

Naturalism, ii, 142 

Naturalist, and evolution theory, 
i, 76, 286; ii, 224; zoological, 
227, 278 

Nature, of things, i, 34; not sim- 
ple, 236; of knowledge, ii, 152 

Nautilus, comparison of, with 
rhizopod, i, 237 

Needs, nutritional, ii, 268 

Neglect nothing, naturalist's mot- 
to, i, 94; ii, 216, 245, 283 

Nerrtertean, regeneration of, i, 
189; capture of prey by, ii, 255 

Neoformation, i, 276 

Nereis, ii, 192 

Neresheimer, E. R., i, 245. 

Nerve, centers, ii, 187; physiology 
of, 320 

Nervous system, in protozoa, i, 
243; autonomic, ii, 128; and in- 
ternal secretions, 128; integra- 
tive action of, 162 et seq.; 
Loeb's important conception of, 

Neural integration, ii, 94, 161 

Neurones, ii, 216 

Neurophanes, i, 245 

Neuro-motor apparatus, i, 243; 
system, 255 

Nickel, ii, 327, 329 

Night-hawk, ii, 258 

Nomenclature of germ-plasm the- 
ory, ii, 87 

Non-cellular, protozoa interpreted 
as, i, 290 

Nucleus, i, 207, 331; in bacteria, 
261; part in pigment formation, 
340; in oxidizing action, 341; of 
"utmost theoretical importance," 
ii, 22; control of cytoplasm by. 


Nusbauin, J., on tissue trans- 
formation, i, 190 

Nusbauin and Oxner, on regenera- 
tion of nemertean, i, 189 

Natural history, special ability of, Objective, side of psychical asso- 
i, 113; methods of studying the ciation, ii, 230; and' "outer," 292 



Observation, method of, ii, 278 

Odors of animals and plants, i, 84 

CEdema, ii, 116 

Oil-drops and globules, in eggs, i, 
213, 215 

Oliver, J. R., ii, 7 

Omnipotence of chromatin in 
heredity, ii, 14 

One-cell stage of organism's life, 
i, 214 

Ontogeny, misuse of term, i, 271 ; 
of protozoa, i, 277; effort to ex- 
plain on elementalistic princi- 
ples, ii, 158 

Optic nerve, ii, 164 

Organ, i, 245 

Organ-forming substances, in the 
ovum, ii, 16; theory of, 141 

Organ-germs, i, 209 

Organelles, i, 248 

Organic formation, and isolated 
fragments, i, 176 

Organic matter, i, 114 

Organicists, i, 7 

Organ-independence, i, 40 

Organism, as a whole, familiarity 
of expression, i, 1; definition of, 
18; distrust of by biologists, 25; 
and its chemistry, 75; and its 
protoplasm, 120; and its cells, 
150; in interpreting the cell, 156; 
substitution of, for cell, 194; 
consisting of one cell, 227; fic- 
tion of structureless, 256; as 
cause, 276; "organless," 232, 
320; hypothetical primitive, 319; 
fundamentally dynamic, ii, 134; 
"body" and "soul" combined, 
151; as causal explanation, 199; 
as cause of chemical transfer- 
mation, 205; a natural object, 
207; attributes and acts of, 215; 
living, 275 ; physico-chemical 
conception of, and conscious- 
ness, 324 

Organism-transforming action of 
thyroid substance, i, 145 

Organismal theory, i, 2, 280; con- 
structive side of discussion of, 
ii, 91; and C. M. Child's results, 
111; and elementalist stand- 
point, 148; in relation to trop- 

isms, 188; of consciousness, 282 

Organismalism, i, 2; and correla- 
tion, 17; remarks on the term, 

Organismalist, ii, 149 

Organization, law of embryonic 
development, i, 14; of infusoria, 
282; of chromosomes, ii, 28; 
physical, and instinct, 310; and 
emotion, 316 

Organizing power of living beings, 
i, 211 

"Organless organisms," i, 232, 320 

Organogenesis, i, 325 

Organoids, i, 248 

Organs, fallacious teachings about, 
i, 242; rudimentary, 277; "can 
belong only to multicellular or- 
ganisms," 281 

Origin of species, i, 4 

Osborn, H. F., i, 320 

Outer layer, universality in or- 
ganisms, i, 301 

Ovum, i, 210; as entity and as 
germinal entity, ii, 15 

Oxidation, in animal body, i, 340; 
ii, 346 

Oxygen, "doscarecious" powers of, 
ii, 204; in relation to conscious- 
ness, 290 et seq.; as respiratory 
substance, 301 ; latent attribute 
of, 341 ; hereditary, ontogenic 
and individual, 347; "activation" 
of, 348 

Pacific Ocean, journey of fur 
seals in, ii, 211 

Pain, ii, 183 

Pairing, promiscuity in, ii, 267 

Paleobotanists, ii, 58 

Paleontologists, i, 322 

Palms, ii, 299 

Pancreas and pancreatic juice, ii, 

Pangens of Darwin, i, 19 

Parallelism, psycho-physical, irre- 
solvable inconsistency of, with 
organismal standpoint, ii, 150, 
220; historical basis of, 297 

Paramecium, i, 326; ii, 253 

Paratoid gland, of toad, i, 111 

Parthenogenetic eggs, i, 351 



Particles, ultimate, ii, 151 

Partisanship in science, i, 338 

Payne, F., ii, 34 

Pearl, R., i, 311 

Peckhams, G. W. and E. S., ii, 

Percepts, ii, 294 

Periodic law, in chemistry, ii, 329 

Personal conscience, ii, 292 

Personality, ii, 335; and person, 
295; and elementary substances, 

Petrels, i, 85 

Phases, fluid, of cell system, i, 
216; of the cell, ii, 326 

Philosopher's stone, ii, 288 

Philosophy, cartesian, ii, 297 

Phleps, Ed., ii, 135 

Phlogiston, i, 225 

Phosphorus, ii, 290; a simple, 287; 
glow of, 343 

Phylogeny, of biochemical sub- 
stances, i, 110 

Phylloxerans, eggs of, i, 352 

Physical basis of life (Huxley), i, 
121; of heredity, ii, 64 

Physical chemistry, in biology, i, 
114; and protoplasm doctrine, 
140; and the organismal stand- 
point, 191; conception of cell, 
215, 333; conception of organ- 
ism, ii, 71; limitations of, in 
biology, 208; absence of in 
earlier biology, 336; anti-ele- 
mentalistic tendency of, 341 

Physical science, ii, 152 

Physio-chemical substances and 
forces, in behavior, ii, 312 

Physics, and chemistry in hered- 
ity, i, 115; province of, 141 

Physiologist, ii, 285 

Physiology, and heredity of mus- 
cle fibers, ii, 61; distinctive task 
of, 274 

Phytin, ii, 123 

Pigeons, control of sex in, ii, 78; 
Whitman's study of habits of, 

Pigment, from chromatin, i, 338; 
bearers, 340 

Pigmentation, i, 213 
Pill bugs, i, 219 

Pillsbury, W. B., ii, 231 

Pineal body, ii, 114 

Pituitary Body, ii, 114, 124 

Plankton, ii, 280 

Plantagens, i, 36 

Planaria, ii, 109 

Plasma, i, 134 

Plasmic elementalism, i, 311 

Play of animals, ii, 273 

Pleasure and Pain, criterion of, i, 

Pleomorphism, i, 266 

Plover, Golden, ii, 259 

Pluralism, philosophical, ii, 294 

Pluteus larva, i, 203 

Poisoning, strychnine, ii, 173 

Polarity, in plants and animals, i, 

Pollen grains, in fertilization, i, 
343; structure of membrane of, 
ii, 57 

Porospora yigantea, i, 270 

Post-generation, i, 207 

Postulate, ii, 297 

Powers, secret, of substances, ii, 

Precipitin reaction, i, 100 

Predisposition, i, 206 

Primitive, i, 290 

Primrose, evening, i, 353 

Primiin movens, i, 276 

Principle, of unity of organiza- 
tion, i, 166; of aggregation, 182 

Principles, Alchemists', ii, 288 

Private opinion, ii, 292 

Promiscuity in mating, ii, 267 

Promorphology, of germ cells, i, 
211; metaphysical, 225 

Properties of the whole, i, 13, 148 

Prophysiology, i, 212 

Protista, i, 230, 280 

Protophyta, i, 230 

Protoplasm, as goal of biology, i, 
5; mystification of, 120; and the 
organism, 120; Max Schultze on, 
125; latest views as to morphol- 
ogy of, 133; specificity of, 143; 
as general term must lie used in 
plural number, 148 

Protozoa, theoretical simplicity of, 
i, 230, 286; development of/2<>7; 
theoretical aggregation of to 



produce metazoa, 268 

Protozoan colony, theory of, as 
nature of metazoan, i, 222 

Protozoology, i, 280 

Pseudopodia, i, 240; pseudopodial 
system, 239 

Psychic activities, subrational, 
four certainties about, ii, 250 

Psychic integration, ii, 94; discus- 
sion of, 214 et seq. . 

Psychic life, subrational moiety 
of, ii, 246; phases of, 274; spec- 
ificity of, 276; man's higher, 
283; catholicity of attitude 
toward, 284 

Psychical, organic connection be- 
tween physical and, ii, 239 

Psychical element, an abstraction, 
ii, 235 

Psycho-analysis, ii, 350 

Psycho-physical parallelism (see 

Psychoids, ii, 149 

Psychologist, ii, 285 

Psychology, associationist, ii, 228, 
and Wm. James, 229; without a 
soul and without a body, 322; 
social and domestic, 332 

Purpose, of reflex, ii, 184 

Python, proportionality of parts 
in skeleton of, ii, 96 

Radicals, compound, in chemistry, 

ii, 343 
Radiolarian, compared with jelly 

fish, i, 235; swarm spores of, 278 
Rage, ii, 183 

Random movements, ii, 252 
Reaction, chemical and neural, ii, 

Reason, ii, 216; "forms the 

world," 243 
Receptors, superficial and deep, 

ii, 173 
Redwood, adventitious buds in, i, 

38; meristic growth series in, ii, 

Reed, H. S., on adventitious buds 

in lemon trees, i, 38 
Reflex arc, cellular integration in, 

ii, 163 
Reflexes, simple, an abstract con- 

ception, ii, 168; compounding 
and spreading of, 171; scratch, 
allied, proprio-ceptive, 172; an- 
tagonistic, 174, 176; inhibition 
of, and compensatory, 177; pur- 
pose of, 184; relation to in- 
stinct, 188, 246 

Regulation, formative, i, 199 

Reichert, E. T., i, 95 

Relationship, the problem of 
causal, ii, 224 

Rennet, i, 104 

Repetition, in organic growth, ii, 
95 et seq.; in instinctive activ- 
ity, 257 et seq. 

Repetitive parts, ii, 95 

Reproduction, asexual, and hered- 
ity, i, 309 et seq. 

Researches in biology, field, labor- 
atory, quantitative, ii, 278 

Resemblance, importance in doc- 
trines of heredity, i, 312, 315; 
and difference, 317 

Respiration, life, and conscious- 
ness, ii, 286 et seq.; of muscle, 

Responses, ii, 216 

"Restlessness," mental, ii, 225, 243, 

Retzius, G., comparative re- 
searches on spermatozoa, i, 216, 
ii, 2 

Reversibility, of direction in nerve 
conduction, ii, 166 

Rhizopod, i, 123 

Rhus, chemistry of, i, 87 

Riddle, O., ii, 76 

Ritter and Forsyth, ii, 45 

Robertson, T. B., on chemical ac- 
tion in growth, ii, 105; on 
tethelin, 123 

Rousseau, J. J., ii, 224 

Roux, W., terminology of "De- 
velopmental mechanics," i, 18; 
Mosaic theory of, 198 

Royce, J., relation between trop- 
isms and apperception, i, 23, ii, 
220; on mental initiative and 
restlessness, 243 

Ruzicka, V., i, 262 

Sachs, J., i, 16; law of, 219 



"San Diego region" (oceanic), ii, 

Saint-Hilaire, J., i, 7 

Sajous, E. de M., ii, 124 

Salamander, ii, 311 

Salmon, ii, 250 

Salpa, i, 316 

Sarcode, identification with plant 
protoplasm, i, 123 

Schafer, E., on endocrine organs, 
ii, 14; drug-like action of inter- 
nal secretion, 122 

Schaudinn, F., i, 261 

Schleiden, M. J., conception of 
plant, i, 34 

Schultze, Max, on protoplasm and 
cell, i, 125 et seq. 

Schwa nn, Th., theory of cells, i, 

Science, positive, i, 298; creative 
impulse in, ii, 227 

Scientific, spirit, i, 338; attitude, 
difference between, and philo- 
sophic, ii, 307 

Scratch-reflex, workings of, ii, 
172; possible "ultimate explan- 
ation" of, 202 

Scripps Institution, statistical 
methods in, ii, 280; for Biolog- 
ical Research, 332 

Sea Urchin, properties of eggs of, 
i, 107; experiments on develop- 
ing eggs of, 202 

"Seat" of heredity, i, 321; of in- 
heritance material, ii, 23; brain 
supposed seat of coordination, 
ii, 191 

"Secret powers" of substances, ii, 

Secretin, ii, 120 

Secretions, internal, integrative 
office of, ii, 113 et seq. 

Secretory systems, internal, ii, 128 

Sedgwick, A., ii, 50 

Seeds, gradations in, i, 102 

Segmental theory of nerve action, 
ii, 185; of nervous system, 190 

Segregation, in heredity, i, 355 

Selection, sexual, ii, 262 

Self-preservation and regulation, 
i, 18; -differentiation, 199, 208; 
ii, 245; -activity, 226; -exhaust- 

ing ceremonies, 264; -injury 
through sex impulse, 267; nat- 
ural history method in study of, 
282; -defence and -preservation, 
292; or person, 301; -control, 
305; -development, 305 

Sellars, R. W., individuality in 
percepts, ii, 294; the individual, 
and social psychology, 332 

Sensationalism, ii, 218 

Sense organ, ii, 165; senses, 216 

Sequoia sempervirens, adventi- 
tious buds of, i, 38; growth 
series in, ii, 101 

Series, graded repetitive, ii, 95 

Serum, rabbit, i, 101 

Sex, cell production, theory of, i, 
61; as unit-character, 348; de- 
pendence upon chromosomes, 
346, 350; impulse, excessiveness 
of, ii, 265; power of, 267 

Sexes, numerical proportion of, ii, 

Shakespeare, Wm., "reckless volu- 
bility" of, ii, 223 

Sherrington, S. C., fundamental 
work on nervous system, i, 22, 
ii, 162 et seq.; neglect of, by 
J. Loeb, 185 

Shrikes, ii, 270 

Simple reflex, an abstract concep- 
tion, ii, 168 

Simplicity, "ultimate," i, 320 

Skin, universal presence of, in or- 
ganisms, i, 300 

Smallpox, i, 264 

Smell, i, 84 

Smith, J. B., i, 214 

Society, and individual, ii, 332 

Soma, i, 319 

Song habits of birds, ii, 260 

Soul, composed of "seminal 
atoms," i, 3; as aspect of organ- 
ism, ii, 215; and body, interac- 
tion between, 323 

Specialist, the unpoised, ii, 247 

Species, difference in egg of, ii, 20 

Specific differences between germ 
cells, importance of, i, 214 

Specification of organic matter, i, 

Specificity, chemical, of organ- 



isms, i, 83; of corresponding 
proteins, 95; of protoplasms, 
143; of sperm of anthropoids 
and man, ii, 2; of animal be- 
havior, 276; of psychical and 
reactive life, 281 

Speculations, i, 293 

Spermatozoa, species diiferences 
in, i, 216; structure of tail of, 
333; structure of head of, 342, 
and ii, 87; two kinds of, i, 347; 
subject to heredity, 399; resem- 
blance to tadpole, ii, 3; ontog- 
eny of mammalian, 4 

Spicules of sponges, ii, 50 

Spinules of Ascidian, ii, 44 

Spirits, historical relation to es- 
sences, ii, 288; historical relation 
to breath and air, 338 

Sponge tissues, i, 144 

Spontaneous, generation, ii, 316; 
in mental activity, 226; on 
meaning as applied to "origin 
of life" (see glossary) 

Spores, i, 269; of mosses and 
ferns, ii, 258 

Squid, i, 221 

Squirrel, storing habits of, ii, 271 

Stage, one celled, of animal life, 
i, 214 

Star fishes, ii, 96 

Starch grains, i, 214 

Stein, Fr., i, 286 

Stentor, supposed nerves in, i, 245; 
ontogeny of, 272 

Stephens, Frank, on storing habits 
of bees, ii, 268; of antelope 
ground squirrel, 271 

Stevens, Miss M. M., i, 350 

Stimuli, summation of, ii, 167 

Storing habits, of honey bee, ii, 
268; of woodpecker, 269; of 
mammals, 270 

Stout, G. F., ii, 298; unanalyzed 
cognition in consciousness, 308; 
on personality in animals, 328 

wStrassburger, Ed., i, 342; and 
chromosome dogma, ii, 259 

Striated muscle tissue and hered- 
ity, ii, 59 

Structurelessness, i, 285 

Structures, ii, 199; constitutively 

antagonistic, 134; organ-form- 
ing, 141 

Struggle, of the parts, ii, 175; for 
existence, 225 

Studies, practical, i, 214 

Sub-conscious, ii, 350 

Subjective, ii, 231; side of asso- 
ciation, 230; and "inner," 292 

Substance, of soul, i, 4; organic 
production of, 81; simple homo- 
geneous, 281; imaginary, 240; 
nutritive, 244; criterion of ele- 
mentary, 287; abuse of the 
term, 298; respiratory, 301; and 
energies, 337; free and vague 
appeals to, ii, 239 

Sugar, and enzyme action, ii, 81 ; 
from liver into blood, 132 

Summation, of stimuli, ii, 167 

Sumner, F. B., ii, 258 

Supernaturalism, and finalism, ii, 
142; and materialism, 148 

Suprarenal body, ii, 114, 135; in 
alliance with thyroid and pitui- 
tary, 127 

Surface energy, of muscle fibers, 
ii, 63 

Surfaces of separation, ii, 170 

Surf perches, matings, habits of, 
ii, 265 

Surgeons, Swiss, ii, 115 

Surplus energy, ii, 273 

Sutton, W. S., i, 356 

Swarm spores, of radiolaria, i, 278 

Swezy, olive, i, 291 

Symbiotic theory of organism, i, 
35, 183 

Symmetry of animals, ii, 189 

Sympathetic nervous sj^stem, ii, 

Synthesis, organizing, ii, 149; as- 
similative, 205; and analysis, 
235; apperceptive, 237 

System of nature, i, 4 

Systematic biochemistry, i, 95 

Tadpoles, of frogs and toads, ii, 


Takamine, ,T., ii, 123 
Taste, i, 84 
Taxonomic, in organic grades, ii, 

42; discrimination, 62 



Taxonomist, i, 213 

Terms, of oxygen, ii, 202; of life, 

Tetany, i, 117, 135 

Tethelin, ii, 123 

Theory, of structive of proto- 
plasm, i, 138; wrong better than 
none, 292; Leibnitzian, ii, 150; 
of organisms, and of knowledge, 
152; elementalist, 157; tropistic 
and segmental, of nerve action, 
185; of tropisms, 232; of apper- 
ception, 232; of knowledge, 296; 
of consciousness, 296 

Thomson, J. A., i, 309 

Threshold of excitability, ii, 165 

Thymus, ii, 114 

Thyroid, ii, 124; apparatus, 114; 
effect of removing, 115, 117; 
triple alliance with pituitary 
and adrenal, 127; substance, 
143; organ-transforming sub- 
stance from, 145 

Tissue, laboratories, i, 83; mix- 
tures, 143; cultures, 168; falla- 
cious teaching about, 240; of 
multicellular organisms, 281 ; 
isolated, 294; of trees, specific- 
ity of, ii, 58 

Toads, ii, 76 

Tonniges, C., i, 326 

Torrey, H. B., ii, 97 

Totipotence, theory of, i, 202 

Transformation, essence of evolu- 
tion, i, 41; heredity works by, i, 
312, 322; of substances, ii, 72, 

Transmission, in connection with 
heredity, i, 312 

Transmutation of metals, ii, 288 

Transcendentalism, ii, 233 

Treviranus, i, 5 

Trial and error, ii, 252, 256 

Trichocysts, i, 326 

Tripylea, i, 278 

Tropism theory, essentially an or- 
ganismal theory, ii, 188; relation 
to apperception, 232 

Tropisms, explained by organism, 
ii, 190; organismal nature of, 
239 ; automatic and anticipatory 
character of, 241 ; higher ra- 

tional life and, 242 

Tropistic and higher psychic ac- 
tivity, ii, 220 

Tropistic mechanism, ii, 189 

Tropistic theories of nerve action, 
ii, 185 

Truth, ultimate, ii, 152, 289 

Trypsin, i, 106 

Type, i, 7 

Tyranny, of the whole, ii, 158 

Ultimate, problem, i, 35; criterior 
of, ii, 149, 201 ; particles, ii, 151 : 
truth, 152, 175, 289; elements. 

Ultramicroscopic organisms, i, 265 

Unicellular, i, 290 

Units, representative, i, 306 

Unity, the organism the "only 
real," i, 12, 26, 205; physiolog- 
ical, 14; of the individual, 33; 
thought of, ii, 150 

Uranium H., ii, 341 

Utility, racial, ii, 251 ; natural se- 
lectionist meaning of, 261 

Variation, ii, 245 

Vertebrates, man a, because moth- 
ers were, ii, 43 

Verworn, M., ii, 279 

Vetch, ii, 106 

Virgin propagation, i, 352 

Vital force, ii, 149 

Vitalism, i, 113; author's attitude 
toward, ii, 207 

Viviparous bony fishes, mating 
habits of, ii, 265 

Volition, ii, 161 

Wallace, A. R., ii, 278 

Waller, H. E., ii, 126 

Warblers, ii, 264 

Wasps, instincts, variability of, ii, 
251 ; excessive action of, 268 

Wassermann, i, 258 

Watase, S., i, 221 

Webber, H. J., i, 330 

Weismann, A., studies of sex-cells 
in hydroids, i, 60; Grette's re- 
sults contrary to, 68; metaphys- 
ics of, 225, 348 

Whale, breaching of, ii, 257 

Wheeler, W. M., sense of smell, 



and odors, of ants, i, 87, 89; 
early embryology of insects, 
218; as field naturalist of mod- 
ern type, ii, 278; on problem of 
instinct, 284; on instinct and 
bodily organization, 311 

Whitman, C. O., as pre-organis- 
malist, i, 11; on cell-theory, 220; 
relation between instinct and 
structure, 314 

Whole, embryo, i, 204; "tyrannizes 
over parts," ii, 159 

Will, feeling, and intellect, ii, 216, 

Wilson, E. B., as pre-organismal- 
ist, i, 11; on cell structure, 135; 
statement of cell-theory, 151 ; on 
"real unity," 192; on early em- 
bryology of amphioxus, 204; on 
promorphology, 217; on x and y 
chromosomes, 350; on connec- 
tion between chromosomes and 
Mendelian inheritance, 356; pro- 
posal to drop "determiner" as 
genetic term, ii, 82; on germ as 
detached portion of parent, 88 

Wilson, H. V., i, 144 

Winterstein, H., ii, 62 

Woodpecker, California, storing 
habits of, ii, 269 

Wood-tissues, ii, 58 

Work, energy, power, force, ii, 342 

World, external, ii, 303 

Wundt, W., and apperception, i, 
23; definition of apperception, 
ii, 233; seeming transcendental 
element in the apperception of, 

Yerkes, R. M., on combined ex- 
perimental and field research in 
behavior, ii, 279 

Ziegler, C., i, 201 
Zoja, R., i, 204 
Zoological diagnosis, i, 265 
Zoologist, anthropological, ii, 285 
Zoology, instruction in elementary, 
i, 236; and the science of be- 
havior, ii, 208; fundamental 
terms of, 247; taxonomic, 276 
Zygote, i, 269